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Gilbert Dennison Harris
(1864 - 1952)

Founder of the Bulletins of American Paleontology (

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SBN 0-87710-4
: SEIS HISS

LUME 100, NUMBER 337

Late Triassic cyrtinoid spiriferinacean brachiopods

from western North America and their

biostratigraphic and biogeographic implications

by

Peter R. Hoover

Paleontological Research Institution
1259 Trumansburg Road
Ithaca, New York, 14850 U.S.A.

OCTOBER 16, 1991 |

CAS hor

PALEONTOLOGICAL RESEARCH INSTITUTIC YN

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OCTOBER 16, 1991

VOLUME 100, NUMBER 337

Late Triassic cyrtinoid spiriferinacean brachiopods

from western North America and their

biostratigraphic and biogeographic implications

by

Peter R. Hoover

Paleontological Research Institution
1259 Trumansburg Road
Ithaca, New York, 14850 U.S.A.

LIST OF ILLUSTRATIONS

Text-figure Page
1. Sketch map showing location of North American collecting localities cited in this report .... 2.0.0.0... 0c c cece eee eee eee eee 64, 65
2. Views) of theicooperculum of Spondylospira lewesensis (Lees, 1934) sec. - joc cee coe eee cee aee sees sees snes eee 76
3. Dorsoposterolateral view of ventral valve of Dagyspirifer fascicostata, n. gen. and n. sp., showing one of the two small, paired, ovate

pedicle foramena that pierce the apices of both the ridged portion of the interarea and the dental plates ....................... idl
4. Anterodorsolateral (interior) view of ventral valve of Dagyspirifer fascicostata, n. gen. and n. sp., showing the horizontal bar that

connects: thedental! plates ‘and lateral'shell wall’ ~...0. se.ace «oc eee one oe ee eo aise cess eke Gert ee ee 78
5. Posterodorsolateral views of portions of a dorsal valve of Phenacozugmayerella mimuncinata, n. gen. and n. sp., showing surface

IMICHOOTMAMENUS 15 scree zsepsoc ck eos ech neucien Biers eeocah acess eats ea ves Dadol Oe ee op Te ny Re ee 91

LIST OF TABLES

Table Page

le Wi pper TnassiciSeries; Stages; substages, and Zones, -52-.4- 50 een sess sere eee sles eee eee ee eee 66
2. Measurements of type specimens of Dagyspirifer fascicostata, n. gen. and N. SP... 2... ee ee ee ce ee cece eee eee eee ee 78
3. Measurements of type specimens of Pseudospondylospira perplexa, n. gen. and N. SP. 2... ee ev vv cece eee eevee eee ee 80
4. Measurements of specimens of Spondylospira lewesensis (Lees, 1934) .... 2.220. ec cece cece ec cc cece ccencccusccvcseresueeeue 82
5. Measurements of the holotype of Spondylospira parmata Hoover, 1983 ............00cccccceccceccueceucccucceuseeceeusceus 84
6: Measurements ofispecimens of Spondylospira tricostaynsisp) ..- 4244s cee so aden eee 85
7. Measurements ofitype’specimens' of Vitimetula parva, n. gen. and\n: sp; ..........+2-os06 sec eee eee eee 87
8. Measurements of type specimens of Zugmayerella americana, N. SP. 2... 2... cece cc nce e ence e ce ceceeceeeuceuseeeeees 88
Ov Measurements of:specimens of. Zugmayerella? spA\ c....-- saaec seen aeac ee neseesee seis te ene nen eee 89
10. Measurements of type specimens of Phenacozugmayerella mimuncinata, n. gen. and N. SP. ..... 2. eee ee ev ee ee eee eee eee eee 90

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS
FROM WESTERN NORTH AMERICA
AND THEIR BIOSTRATIGRAPHIC AND BIOGEOGRAPHIC IMPLICATIONS

By
PETER R. HOOVER

Paleontological Research Institution
1259 Trumansburg Road
Ithaca, NY 14850-1398, U.S.A.

ABSTRACT

Cyrtinoid spiriferinacean brachiopods in samples from over fifty Late Triassic marine fossil localities in western North America
include examples of three species of Spondylospira Cooper, 1942, one of which (S. tricosta) is new; one new species of Zugmayerella
Dagys, 1963; and four new monotypic genera, represented by Dagyspirifer fascicostata, Phenacozugmayerella mimunicinata,
Pseudospondylospira perplexa, and Vitimetula parva, all new species. In North America, all of these genera are limited to the
Karnian and/or Norian. One species, Spondylospira tricosta, n. sp. appears to be a good indicator of the early Norian. The family
Laballidae of earlier workers is divided into the family Laballidae and the new family Spondylospiridae, based on absence or
presence, respectively, of functional pedicle foramena; the latter family is divided into two new subfamilies, Spondylospirinae
and Dagyspiriferinae, based on the form of those foramena.

Two features of cyrtinoid spiriferinacean species biogeography have tectonostratigraphic implications. First, there are no
“Tethyan” species among them, and second, the nine species of cyrtinoid spiriferinaceans reported here are predominantly
confined to the Americas, including both cratonal and terrane belt localities. Biogeographic evidence from the cyrtinoids, therefore,
does not support close links between terranes of the present-day eastern and western Pacific.

Cyrtinoid spiriferinaceans provide abundant evidence of biogeographic “‘stitching” of various terranes. For example, Pseu-
dospondylospira perplexa, Spondylospira lewesensis, and Spondylospira tricosta occur in localities that are craton-bound as well

as in parautochthonous and allochthonous localities.

ACKNOWLEDGEMENTS

I owe a great debt of gratitude to Dr. Cathryn R.
Newton (Syracuse University, Syracuse, NY) for her
help with the tectonostratigraphic summaries of the
various localities, and for discussions of the paleogeo-
graphic implications of this fauna. I am indebted to
the National Research Council for awarding me a post-
doctoral fellowship at the United States Geological
Survey’s Paleontology and Stratigraphy Branch at the
United States National Museum of Natural History in
Washington, DC, during which this study was initi-
ated. I am also indebted to the Trustees of the Pale-
ontological Research Institution, of Ithaca, New York
for encouraging me to take the necessary time to com-
plete the study.

Drs. J. Thomas Dutro, Jr. (U. S. Geological Survey,
Washington, DC, and my advisor in the post-doctoral
study which led to this paper), Norman J. Silberling,
(U.S. Geological Survey, Denver, CO), Algirdas S.
Dagys (Institute of Ecology, Vilnius, Lithuania), and
Derek Ager (Swansea, Wales) reviewed the manuscript
and made valuable comments, many of which have
been incorporated into the manuscript.

Iam grateful to Fred Collier (U. S. National Museum
of Natural History, Washington, DC, U. S.A.), Dutro,
Silberling, George Stanley (University of Montana,
Missoula, MT, U.S. A.), and E. T. Tozer (Geological

Survey of Canada, Vancouver, British Columbia,
CANADA) for the opportunity to examine collections
under their care, and for loans of specimens, encour-
agement, constructive criticism, and gracious hospi-
tality during the course of this study.

The illustrations in this study came from a variety
of sources. The photographs on all the plates were
produced in 1975 by R. H. McKinney and H. E. Moch-
izuki of the U. S. Geological Survey’s Paleontology and
Stratigraphy Branch in Washington, DC. I drafted Text-
figure 1; the photographs in Text-figure 2A and B were
shot by McKinney and Mochizuki, and Jorge Valdes,
of S. T. Pees and Associates, Meadville, PA, respec-
tively; Text-figures 3 and 4 come from the talented pen
of Jude Louviere of Toledo, OH; and the photographs
that comprise Text-figure 5 were expertly shot and
printed by David Tuttle, of the Department of Geo-
logical Sciences at the State University of New York
at Binghamton.

INTRODUCTION

Late Triassic cyrtinoid spiriferinacean brachiopods
from western North America historically have been
poorly known. This is partially due to their rarity (com-
pared with other Late Triassic invertebrate fossils) and
partially due to the comparatively greater biostrati-
graphic utility of other forms (e.g., ammonites and flat
clams, and, more recently, conodonts; see discussion

64 BULLETIN 337

Li
AN aN
‘. x Nee ae ny
a cae x
. eee .. jo ee
x va
Q :
< .
U2 . Koq-31
13,17 oe ¢ Se aoe
ae cee aU o> oo 6
53 20° fo & Secs
° ; ; TOS
= uf
; ost
a
0?

Text-figure 1.—Sketch map showing location of North American collecting localities cited in this report (arrows point North).

below). Although there were sporadic reports under
the guise of Cyrtina Davidson, 1858 and Spiriferina
d’Orbigny, 1847 during the late nineteenth and early
twentieth centuries, not until Cooper’s work of 1942
and 1944, which defined Spondylospira, were these
forms studied in appropriate detail. In 1944, in Shimer
and Shrock’s Index Fossils of North America, Cooper
illustrated a specimen he assigned to Psioidea sp. (p.
359, pl. 140, figs. 52-55). This specimen was differ-
ted from the species of Spondylospira illustrated

by having a smooth fold and sulcus. The il-

shows a very smooth surface, and exami-

the specimen (USNM 106250) reveals it to

he internal mold of a cyrtinoid spiriferinacean of

a size and form similar to Spondylospira lewesensis
(Lees, 1934). Because it lacks any remaining vestige of
the shell, it cannot confidently be assigned to any known
cyrtinoid spiriferinacean genus or species. These stud-

ies were followed in short order by the Vokes and Haas
(1944) compendium of information on American spe-
cies of Spondylospira, which incorporated available
information from the Triassic of Peru (KOrner, 1937;
Tilmann, 1917). Since then most North American Late
Triassic cyrtinoid spiriferinacean brachiopods have
been referred to Spondylospira, including some forms
that were not even spiriferinaceans (e.g., Spondylospira
alia Hall and Whitfield, 1877; see discussion under
Spondylospira lewesensis). In recent years, some cyr-
tinoid forms that lack a costate fold and sulcus have
been assigned to Zugmayerella Dagys, 1963 [see e.g.,
Zugmayerella uncinata (Schafhautl, 1851), cited in
Stanley, 1979, p. 14, pl. 8, figs. 10-13], but none has
been formally described.

In 1976, as part of a post-doctoral project, I began
a study of Late Triassic articulate brachiopods, based
primarily on materials deposited in the stratigraphic

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER

s
N
14 aN
a
.
4 ‘

65

~ 15

> Oi

47 fy a12 ,

é 28 ’

; oi a hee
ie Sy ~~

46, ve =
sB:49: at ’ 32,41,43 aS

ee 18,39, stig a

Text-figure 1.—[continued].

collections of the U.S. Geological Survey in Wash-
ington, DC, and Menlo Park, CA. This eventually in-
volved over 50 collecting localities, spread over an area
from central Nevada north to southwestern Alaska
(Text-fig. 1). It became immediately apparent that time
and equipment were inadequate to do justice to the
terebratulids and rhynchonellids, both of which re-
quire elaborate preparation of many specimens to ad-
equately understand the variation in expression of tax-
onomically significant characters. The spiriferoids were
commonly better preserved, and their usable taxonom-
ic characters more readily accessible. Of these, the cyr-
tinoid spiriferinaceans were a convenient group to work
on, as they contain a variety of structures that are of
great interest in terms of functional interpretations.
Indeed, these puzzling functional interpretations are
one of several important factors that have delayed pub-
lication of this monograph (see Hoover, 1983; 1990).

Fourteen years later, in March of 1990, having be-
come increasingly aware of the many fine Late Triassic
marine fossil localities that had been collected from
by Canadian workers, I visited the Geological Survey
of Canada in Ottawa, and surveyed its Triassic bio-
stratigraphic collections. At the Canadian Survey, as
in the U.S., fossils are most commonly collected by
workers engaged in mapping, and chiefly interested in
those forms that have biostratigraphic significance. Be-
cause most biostratigraphically significant Triassic in-
vertebrate macrofossils are not brachiopods, but rather
ammonites or flat clams, there were few brachiopods
in the collections. Indeed, the only Canadian localities
represented in the GSC biostratigraphic collections that
contained cyrtinoid spiriferinaceans were: (1) Nicola,
British Columbia [no further information available],
which yielded a single ventral valve questionably re-
ferable to Pseudospondylospira, n. gen.; (2) the type

66 BULLETIN 337

Table 1.—Upper Triassic Series, Stages, Substages, and Zones (after Tozer, in House and Senior, 1980).

Upper Norian
(Sevatian)(UNo)

Middle Norian

Nomani(Ne) (Alaunian)(MNo)

Lower Norian
(Lacian)(LNo)

Upper Karnian

Karina) (Tuvalian)(UC)

Lower Karnian
(Julian)(LC)

area for Spondylospira lewesensis (Lees, 1934) [Lake
Laberge region, Yukon Territory]; (3) another area in
similar strata about 10 km to the east that also yielded
S. lewesensis [Lime Peak, Yukon Territory (Reid,
1985)]; (4) the Karnian of Cameron Island in the Ca-
nadian Arctic [GSC loc. 25872]; (5) Trout Lake, Yukon
Territory [GSC loc. 86206], which yielded isolated
specimens that may be referable to Zugmayerella Da-
gys, 1963; and (6) the Halfway River area of British
Columbia [GSC loc. 47016], which yielded isolated
specimens questionably referable to the genus Spon-
dylospira Cooper, 1942. This should in no way be con-
strued as a gauge of the rarity of cyrtinoid spiriferi-
naceans in Canada, any more than U.S. collections
are a gauge of their comparative rarity in the U.S.
Rather, they are rarely collected. Because this paper
demonstrates the biostratigraphic utility of some of
these forms, they may be better sampled in the course
of future studies.

Here, then, are six genera of spiriferinacean bra-
chiopods that may prove useful in future paleoecolog-
ical and biostratigraphic studies of the Triassic System.

BIOSTRATIGRAPHY

All specimens discussed herein were recovered from
institutional collections, and their occurrence in the
field was documented by workers whose primary in-
terest was not necessarily biostratigraphic, or even pa-
lecontologic. Thus, much of the information relating to

‘lecting localities themselves is interpretive and was

| from evidence recovered subsequent to the

ing itself (see Appendix, material in brackets).

obable age assignments for the rocks recovered

)calities discussed herein vary in their precision;

yme are made on the basis of stratigraphic position,
and others on the basis of associated fauna.

Dagyspirifer fascicostata, n. gen. and n. sp., and Vi-

iimetula parva, n. gen. and n. sp., each known from a

single locality, occur, respectively, in the earliest No-

rian and late Norian [zones LNol, and UNol through

Choristoceras crickmayi Zone (UNo3)

Cochloceras amoenum Zone (UNo2)

Gnomohalorites cordilleranus Zone (UNo1)

Himavatites columbianus Zone (MNo2)

Drepanites rutherfordi Zone; Cyrtopleurites bicrenatus Zone (MNo1)
Juvavites magnus Zone (LNo3)

Malayites dawsoni Zone; Malayites paulckei Zone (LNo2)
Mojsisovicsites kerri Zone; Guembelites jandianus Zone (LNo1)
Klamathites macrolobatus Zone (UC3)

Tropites welleri Zone; Tropites subbullatus Zone (UC2)

Tropites dilleri Zone (UC1)

Sirenites nanseni Zone (LC3)

Austrotrachyceras austriacum Zone (LC2)

Trachyceras desatoyense Zone; Trachyceras aonoides Zone (LC1)

UNo3 of Tozer (1980); see Table 1]. Their full bio-
stratigraphic ranges are not known.

Pseudospondylospira perplexa, n. gen. and n. sp. oc-
curs in beds that range from early late Karnian to latest
Norian in age [zones UCI through UNo3 of Tozer
(1980)], but most occurrences are in the early Norian.
The precision of dating of these localities is, for the
most part, insufficiently high to make this form a re-
liable biostratigraphic indicator.

Spondylospira Cooper, 1942 has commonly been
used as an indicator for the Late Triassic, and with one
possible exception (Hoover, 1990), that is still true. S.
lewesensis (Lees, 1934), the best known species of the
genus, has been recovered in this study alone from 22
localities ranging in probable age from earliest Karnian
through latest Norian [zones LC] through UNo3 of
Tozer (1980)]. The precision of the age assignments on
the early end of this range is unknown. Only a single
locality (loc. 26, Alexander terrane) includes rocks that
may be older than late Karnian [zone UCI of Tozer
(1980)]. This date is based on analysis of associated
fauna by Silberling (written commun., 1989). Dagys
(written commun., 1990) states that the “‘distribution
{herein] of S. /ewesensis through all Late Triassic is
quite unusual for Triassic cyrtinoid brachiopods. In
Tethys, as well as in Boreal region, they are indicators
only of part of one stage. Very suspect is distribution
of /ewesensis in Carnian.”’ It may be: (1) that the ages
presented here are correct; (2) that age assignments of
some of the localities included herein are too broad or
incorrect; or (3) that some of the material may belong
to other taxa. I believe alternative (3) to be unlikely.
S. parmata Hoover, 1983 is known from only two
localities, but seems such a distinctive form that it
could be a reasonable indicator for the middle to late
Norian [zones MNol through UNo3 of Tozer (1980)].
S. tricosta, n. sp. qualifies as a very good index fossil.
In this study it occurs at six localities in three different
terranes, and nowhere is found earlier than the earliest
Norian nor later than the late early Norian [zones LNol

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 67

through LNo3 of Tozer (1980)]. Because it is morpho-
logically highly distinctive and easily recognizable, it
should find great utility as an index for the early No-
rian.

Zugmayerella Dagys, 1963 as a genus does not ap-
pear to occur earlier than the early late Karnian [zone
UCI of Tozer (1980)] nor later than the late middle
Norian [zone MNo2 of Tozer (1980)], and there are
interesting patterns within the species recognized here-
in. Z.? sp. A is known from only two specimens, from
two localities, one of which is of unknown age, and the
other of which is dated as middle late Karnian [zone
UC2 of Tozer (1980)] on the basis of associated fauna.
Z. sp. is known from a single specimen recovered from
beds that are dated from early late Karnian through
middle early Norian [zones UC1 through LNo2 of Toz-
er (1980)], based on stratigraphic position. Neither can
be considered as a good index based on this small
information base. Z. americana, n. sp., however, is
known from four localities representing three terranes,
all dated on the basis of associated fauna and covering
the relatively short time range of from middle late
Karnian through early early Norian [zones UC2 through
LNol of Tozer (1980)].

Phenacozugmayerella mimuncinata, n. gen. and n.
sp. is a particularly interesting story. It has a completely
disjunct distribution. Four localities, representing the
Alexander (loc. 6) and Izee (locs. 48, 49, 51) terranes,
give an age spanning the Late Karnian [zones UCI
through UC3 of Tozer (1980)], based on associated
fauna in the former and on stratigraphic position in
the latter, but these ages may not be so well constrained
as represented. Eight other localities, representing the
Paradise and Jungo terranes, and rocks overlying the
Golconda allochthon, give an age mostly in the early
Norian, with one locality each from the Paradise and
Jungo terranes extending the range up through the mid-
dle Norian [through zone MNo2 of Tozer (1980)].

EVOLUTIONARY RELATIONSHIPS OF THE
BRACHIOPODS

All of the brachiopods described herein are repre-
sentatives of the families Spondylospiridae or Labal-
lidae of the Superfamily Spiriferinacea. With the pos-
sible exception of a single species of Spondylospira
Cooper, 1942 from Peru (Hoover, 1990), none is known
from strata older than Karnian or younger than No-
rian. Where these two structurally bizarre spiriferina-
cean families came from is a matter of conjecture. After
many years of pondering this problem I am no nearer
to a solution. How evolution proceeded within the
group is somewhat easier to postulate. The two families
are differentiated largely on the basis of the apparent
presence in the Spondylospiridae of a large functional
pedicle in the adult stage, while this feature is only

rarely reported in the Laballidae. Dagys (1974) has
reported a small pedicle foramen in Thecocyrtella Bitt-
ner, 1892, and in Thecocyrtelloidea Yang and Xu, 1966
(pp. 113, 114, pl. 9, figs. 1-14; pl. 10, figs. 2-3), mul-
tiple pedicle tubules that pierce the deltidium have
been described. Although Yang and Xu state [p. 113]
that Thecocyrtelloidea is typified “by . . . lack of dental
plates (hence no spondylium),” Dagys (written com-
mun., 1990) feels that a true spondylium is present.
Thus it appears that if the descent of mutiple-foramen
cyrtinoid spiriferinaceans is monophyletic, this occur-
rence in the Ladinian of Gueizhou, China is a logical
precursor to Spondylospira and related forms in the
later Triassic.

Dagys (1974, p. 284) demonstrates that the Spiri-
ferinacea, like other groups of Paleozoic brachiopods
that survived the great reorganization near the Perm-
ian—Triassic boundary, continued to diversify through-
out the Triassic. In the Early Triassic (Induan and
Olenekian), the superfamily consisted of only a single
genus; in the Anisian, it had eight genera; in the Lad-
inian, nine; in the Karnian, 19; and in the Norian 12.
Studies since 1974 have increased these numbers, but
the pattern remains the same. Dagys (1974, Table 3,
pp. 287-289) indicated that Zugmayerella had a fossil
record covering the entire Norian, while Spondylospira
was known only in the Upper Norian. From evidence
developed herein, I think it is safe to extend the lower
boundaries of both ranges, because good examples of
the families Spondylospiridae (typified by Spondylos-
pira) and Laballidae (typified herein by Zugmayerella)
exist in the Karnian. Dagys (1974, p. 228) cites the
obvious homeomorphy between Paleozoic Cyrtinidae
(which extend from the Late Silurian through the Early
Carboniferous) and the Late Triassic laballids [= La-
ballidae + Spondylospiridae herein], but points out
that there is no direct phylogenetic connection between
the two lineages. Dagys (1974, p. 228) also discounts
the possibility that punctae developed more than once
in the spiriferids (i.e., that these forms were polyphy-
letic), and believes that the Late Triassic spiriferina-
ceans (including those discussed herein) evolved from
spiriferinacean ancestors, not directly from homeo-
morphically-similar cyrtinids. He further suggests that,
although the jugal structures of Late Paleozoic spiri-
ferinids have been insufficiently studied, one detail of
these forms stands out — the jugum is V-shaped and
has its apex directed toward the anterior margin; that
is, its structure is intermediate between the jugum of
cyrtinids and the jugum of younger spiriferinaceans.

Within the Late Triassic spiriferinaceans that have
ridges on part or all of the interarea, the course of
evolution is clearer (Hoover, 1990). Dagysprifer fas-
cicostata, n. gen. and n. sp. (Pl. 9, figs. 1-9), with its
globose form and small ridged portion of the interarea,

68 BULLETIN 337

appears a likely precursor morphology to that of Spon-
dylospira; it is known only from a single locality in the
late Karnian of southeastern Alaska. Forms like Pseu-
dospondylospira perplexa, n. gen. and n. sp. (Pl. 10,
figs. 14-24) from the late Karnian to late Norian of
Nevada, Oregon, and Alaska, have a slightly less glo-
bose form and relatively larger ridged portion of the
interarea, and they seem likely intermediates in this
lineage. This leads through increased relative height of
the ventral valve, broadening of the “‘interarea’’, in-
creased development of multiple ““honeycomb”’ fora-
mena at the ventral valve apex, and decreasing shell
thickness, to forms like Spondylospira parmata Hoo-
ver, 1983 (Pl. 10, figs. 8-13) in the late Norian.

BIOGEOGRAPHY OF THE BRACHIOPODS
AND IMPLICATIONS FOR
TERRANE COMPARISONS

The nine clearly recognizable species of cyrtinoid
spiriferinaceans occur in a wide array of different tec-
tonic blocks within the western Cordillera. Therefore
they provide a new data set for biogeographic com-
parisons among various Cordilleran terranes. This is
the first such data set published for Early Mesozoic
brachiopods of the Cordillera.

Of the six spiriferinacean genera in two families of
cyrtinoid brachiopods decribed in this paper, only two
(Spondylospira Cooper, 1942, and Zugmayerella Da-
gys, 1963) appear to be known outside North America,
and of these, only Zugmayerella is cosmopolitan. In
addition to North American occurrences, Spondylo-
spirais also known from South America and the north-
easternmost U.S.S.R. (Bychkov and Dagys, 1984;
Dagys, written commun., 1990). The two spondylo-
spirid genera Pseudospondylospira, n. gen. and Viti-
metula, n. gen., are known from but a single locality
each, in southeastern Alaska [zones UCI-—UNol of
Tozer (1980); Alexander terrane] and Idaho [zones
UNol1-UN6o3 of Tozer (1980); Wallowa terrane], re-
spectively. Zugmayerella, when first described by Da-
gys (1963), was reported from the Norian and Rhaetian
of the Alps, the Carpathians, the Crimea, the Caucasus,
northwestern Turkey, and the northeastern U.S. S. R.
(see Bychkov et a/, 1976). It has since been recognized
in Alaska (Wrangellia terrane), California (Eastern

‘amath terrane) and Oregon (Wallowa terrane) [this

Chile (Chong D. and Hillebrandt, 1985), Peru
unpublished data), Papua New Guinea

‘icoll, and Campbell, 1976), China (Ching

ig, 1977), Tibet (Sun, 1981), and New Zealand
impbell, 1990). Stanley’s (1979) report of Zugmay-
rella in the Late Triassic of Nevada was based on my
OWn erroneous preliminary identifications; in the light
of the present report, these forms are probably better
assigned to Phenacozugmayerella, n. gen. Phenaco-

zugmayerella, n. gen., in many respects a homeomorph
of Zugmayerella, is known from 12 localities in Alaska
(Alexander terrane), Nevada (Paradise terrane, Jungo
terrane, and Triassic cover on the Golconda Allo-
chthon), and Oregon (Izee terrane).

Three features of cyrtinoid spiriferinacean species
biogeography are particularly striking and offer insight
into terrane comparisons and paleogeography. First,
true ““Tethyan” species are absent. Indeed, only one of
the six genera reported here (Zugmayerella) includes
species that occur in Tethys, but the closest Tethyan
relative of an American species is from China, at the
eastern end of Tethys. Dagys (written commun., 1990)
suggests that “Zugmayerella is [a] cosmopolitan genus,
but species are different in remoted [sic] regions —
western Tethyan, eastern Tethyan (China), Boreal (Si-
beria), eastern Pacific, and perhaps Notal. North
American species were connected with Tethyan rather
via boreal regions. True Tethyan species of cyrtinoid
brachiopods (as well as other characteristic Tethyan
species of this group — koninkinids, thecospirids, etc.)
were absent in East Pacific region.”

Second, some of the forms endemic to North Amer-
ica occur in several different types of terranes. Phen-
acozugmayerella, n. gen. occurs not only in the al-
lochthonous Alexander, Paradise, and Izee terranes,
but also in parautochthonous terranes such as the Jun-
go, and in rocks associated with blocks such as the
Golconda Allochthon, which were linked with North
America and ‘‘autochthonous” by the Late Triassic.

The distribution of Zugmayerella parallels that of
the Late Triassic rhynchonellid brachiopod Halorella
Bittner, 1884, whose geographic occurrences have been
summarized by Ager (1964). Halorella occurs over a
broad geographic range within the Tethyan seaway,
and also has been recognized both in allochthonous
Cordilleran terranes of eastern Oregon (Ager, 1964),
and in rocks of the parautochthonous Jungo terrane
(Burke and Silberling, 1973).

The nine species of cyrtinoid spiriferinaceans re-
ported herein are mostly confined to North and South
America, including the terrane belt. The only taxo-
nomic overlap with Western Pacific Norian assem-
blages is with the species of Spondylospira from the
Koryak Range of the northeasternmost U.S.S. R.
Cyrtinoid spiriferinacean brachiopods show little bio-
geographic evidence for linkages among terranes of the
present-day eastern and western Pacific.

Abundant evidence does exist for biogeographic
“stitching” among the North American terranes — in-
cluding biogeographic links among allochthonous, par-
autochthonous, and craton-related terranes. Among the
spiriferinaceans, several species illustrate these com-
plex, overlapping distribution patterns particularly well.

(1) Pseudospondylospira perplexa, n. gen. and n. sp.

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 69

is known from the allochthonous Alexander and Wal-
lowa blocks. However, it also occurs in the Triassic
cover on the Golconda Allochthon, which is thought
by most workers to have been tectonically emplaced
against the North American craton during Permian to
Early Triassic time (Miller et a/., 1984; Brueckner and
Snyder, 1985; Silberling et a/., 1987; Babaie and Speed,
1990). New structural and stratigraphic data (Ketner,
1990) indicate that parts of the Golconda Allochthon
overthrust Lower Triassic strata as a result of Mid- or
Late Mesozoic tectonism. However, the youngest time
of initial emplacement is mid- to Late Triassic (Sil-
berling et a/., 1987; see discussion under “Triassic cov-
er on the Golconda Allochthon’’, p. 71). Thus, occur-
rences of P. perplexa in rocks overlying the Golconda
Allochthon represent localities whose paleogeographic
position was either on or very near the Late Triassic
North American continent.

Pseudospondylospira perplexa also occurs in the par-
autochthonous Jungo terrane of Nevada. Triassic stra-
ta of the Jungo terrane have been identified by some
workers as basinal equivalents of the Auld Lang Syne
Group of the Golconda terrane, which was craton-
bound by latest Triassic time (Silberling et a/., 1987).
The depositional cyclicity within the Auld Lang Syne
Group in turn has been correlated with that of the
North American Chinle Formation (Lupe and Silber-
ling, 1985). An alternative recent interpretation by Ol-
dow, Bartel, and Gelber (1990) is that the basinal Early
Mesozoic rocks of the Jungo terrane (““Lovelock as-
semblage” of Oldow, Bartel, and Gelber) are alloch-
thonous relative to the neighboring craton-related plat-
formal sediments of the Golconda terrane. However,
Oldow (1984) has interpreted the Jungo—Lovelock Ear-
ly Mesozoic sediments as having been thrust eastward
during closure of a marginal basin behind the Sierra
Nevada arc, and thus, even this “‘allochthonous”’ in-
terpretation of Jungo strata does not involve large-scale
longitudinal displacement of Jungo rocks. Therefore,
the Jungo terrane is probably parautochthonous. Note
that the occurrence of P. perplexa in the Jungo litho-
tectonic block is significant in bridging the boundary
between the craton-related Golconda terrane and the
more outboard, truly allochthonous Cordilleran blocks
such as the Alexander terrane. In summary, then, P.
perplexa is here reported from occurrences that rep-
resent craton-bound North American rocks, parauto-
chthonous rocks, and allochthonous rocks.

(2) Spondylospira lewesensis (Lees, 1934) is known
from a broad range of allochthonous terranes within
the North American Cordillera, including Eastern
Klamath, Alexander, Paradise, Wrangellia, Wallowa,
and Chulitna. It also occurs in the parautochthonous
Jungo terrane (see discussion above), and is found in
rocks overlying the Golconda allochthon, which by

Late Triassic time was likely bound to the North Amer-
ican craton (see discussion above). In addition, S. /ew-
esensis is known from the Stikine terrane (where it was
first described), which is inferred to have amalgamated
with Cache Creek by Late Triassic time and may have
accreted to North America during Jurassic time (Mon-
ger, Price, and Templeman-Kluit, 1982; Cordey et al.,
1987). Paleomagnetic data from Permian and Triassic
strata of the Stikine block initially were thought to
indicate significant northward latitudinal displacement
of Upper Triassic and younger rocks. Late Triassic
reefs described by Reid (1985) further suggest an origin
for Stikine far south of the terrane’s present position.
However, May and Butler (1986) have shown that when
revised North American Upper Triassic poles are used,
the Stikine Triassic results are in agreement with ex-
pected cratonal inclinations. Comparable results have
been obtained for Permian rocks of the Stikine block
(Irving and Monger, 1987). Thus, paleomagnetic data
show no indication of major post-Permian translati-
tudinal motion for Stikine, leading some workers to
speculate that Stikine may have ‘evolved near, but
seaward of, the margin of North America” (Samson et
al., 1989, p. 705). The presence of S. /ewesensis in the
Stikine block and in the Golconda terrane shows that
this species inhabited North American parauthoch-
thonous and cratonal habitats as well as occurring in
more distant allochthonous terranes. Thus, S. /ewes-
ensis provides yet another biogeographic tie between
cratonal North America and the allochthonous ter-
ranes.

(3) Spondylospira tricosta, n. sp. also occurs in ter-
ranes of diverse origins. It is known from the craton-
bound Triassic strata overlying the Golconda allo-
chthon, but it is also represented in the Paradise terrane
of Nevada and the Wallowa terrane of northeastern
Oregon and Idaho. The distribution of S. tricosta com-
pares with that of Pseudospondylospira perplexa, n.
gen. and n. sp. and S. /ewesensis in that this species
also occurs in both craton-bound (Golconda) and “‘sus-
pect”’ terrane localities.

In summary, the absence of ““Tethyan”’ species and
the complex, overlapping biogeographic patterns of
spiriferinacean brachiopods in this study compare fa-
vorably with patterns previously described for Cor-
dilleran bivalves by Newton (1987). The most signif-
icant pattern documented in this study is the
biogeographic “stitching”? of North American craton-
bound brachiopod faunas with those of the parauto-
chthonous and allochthonous Cordilleran terranes.
These shared brachiopod species suggest that either
many terranes originated close to the North American
continent, or that there was large-scale dispersal of
faunas within the proto-Pacific ocean basin. Evidence
for the latter is also suggested by apparent species co-

70 BULLETIN 337

occurrences in Spondylospira in North America and
in the northeasternmost U.S. S.R., and similarities
between North American and Chinese species of Zug-
mayerella.

TECTONOSTRATIGRAPHIC
SETTING OF LOCALITIES

The following discussions of North American ter-
ranes are ordered geographically, from south to north,
and are followed by a brief discussion of the marine
sedimentary rocks of the Cerro de Pasco region of Peru,
from which Late Triassic cyrtinoid spiriferinacean bra-
chiopods have also been recovered. In these discus-
sions, a terrane is a descriptive unit having no inherent
genetic implications (sensu Dover, 1990).

JUNGO TERRANE
[Churchill and Pershing counties, Nevada]
(locs. 18, 39, 45)

The Jungo terrane of northwestern Nevada preserves
an Early Mesozoic stratigraphy of deformed Upper
Triassic-Lower Jurassic fine clastic strata that repre-
sent basinal facies (Silberling, 1990). This terrane, as
defined by Silberling et a/. (1987) and Silberling (1990),
is the equivalent of the “Lovelock assemblage” dis-
cussed by Oldow (1984) and Oldow, Bartel, and Gelber
(1990). Lupe and Silberling (1985) and Silberling ef al.
(1987) infer that these basinal Jungo strata are the
distal equivalents of the Auld Lang Syne Group sed-
iments that overlie the Golconda Allochthon. Oldow,
Bartel, and Gelber (1990) infer that the Late Mesozoic
Fencemaker thrust, which separates the Early Meso-
zoic rocks of these two terranes, coincides approxi-
mately with the sedimentological transition between
equivalent platformal (Golconda) and basinal (Jungo)
strata. The Upper Triassic sedimentary units of the
Jungo terrane include the lower Norian Hollywood
Formation and overlying Antelope Springs Formation,
the middle Norian Lori Formation, and the upper No-
rian to Lower Jurassic Packard Wash Formation (Ol-
dow, Bartel, and Gelber, 1990).

PARADISE TERRANE
[locs. 8-11, 33-38, 40]
(Mineral and Nye counties, Nevada)

Nevada terrane was initially assigned

k within the Walker Lake terrane of Sil-

: (1987), but recently Silberling (1990) el-

ted the Paradise to terrane status. This terrane, as
scognized by Silberling (1990) and Silberling et al.
(1987), is equivalent to the Sand Springs, Pamlico,
Luning, and Gold Range lithotectonic assemblages of
Oldow (1984). The Paradise terrane consists of Permo-

Triassic volcanogenic rocks, overlain by Upper Tri-
assic to Lower Jurassic platformal carbonates that also
include clastic and volcanogenic components (Silber-
ling, 1990). Upper Triassic formations within the Par-
adise terrane include the Luning and Gabbs forma-
tions, both of which yield cyrtinoid spiriferinacean
brachiopods. The stratigraphy of the Luning—Gabbs
sequence was first described by Muller and Ferguson
(1936; 1939) and, in part, has most recently been re-
viewed by Taylor et a/. (1983). The Luning, principally
of Norian age, represents a carbonate platform with
interspersed clastic sediments; it is overlain by clastic
sediments of the upper Norian Gabbs Formation, which
is overlain in turn by lowermost Jurassic clastics of the
Sunrise Formation (Taylor ef a/., 1983; Silberling et
al., 1987). Much biostratigraphic work has been done
on rocks of the Gabbs and Sunrise formations, because
the Triassic-Jurassic boundary coincides approxi-
mately with the formational contact (Guex and Taylor,
1976; Hallam, 1981; Laws, 1982; Taylor et al., 1983;
Tozer, 1988).

The Paradise terrane is clearly, in the strict sense,
allochthonous with respect to cratonal North America,
because it is bounded by thrust faults (e.g., Silberling
and John, 1989). Exposures of Paradise terrane de-
scribed by Silberling and John (1989) occur west of the
Sr,;=0.706 isopleth, which is interpreted as the west-
ernmost extent of Precambrian continental basement
(Levy and Christie-Blick, 1989, and references there-
in). However, the magnitude of post-Early Mesozoic
displacement relative to the North American continent
is not known, and the Mesozoic rocks of the Paradise
are often interpreted as parautochthonous rather than
allochthonous; for example, Speed, Elison, and Heck
(1988) infer that the Paradise rocks were deposited
atop an allochthonous Paleozoic basement after su-
turing to North America. Some tectonic studies (e.g.,
Silberling, 1990) have also preferred a fringing-arc in-
terpretation of the volcanogenic sediments of this and
related terranes, in which these Nevada terranes orig-
inated close to the American continental margin but
were tectonically reshuffled later in the Mesozoic. One
specific variant on this theme was proposed by Oldow
(1984), who inferred that the thrusting of the Luning—
Fencemaker and other western Nevada allochthons re-
flects closure of a back-arc basin behind (eastward of)
the Sierra Nevada arc. In this last view, although there
may have been hundreds of kilometers of southeast-
directed thrusting, the western Nevada terranes would
essentially be parautochthous with respect to cratonal
North America. More recent work by Oldow, Bartel,
and Gelber (1990) has shown that, although there are
lithostratigraphic differences among the western Ne-
vada terranes (including the Paradise terrane), prov-
enance linkages between stratigraphic units do exist —

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 71

for example, between the platformal Triassic sedi-
ments of the Golconda terrane and correlative sub-
marine fan deposits of the Jungo terrane. Biogeograph-
ic evidence from ammonoid cephalopods supports the
idea that the Triassic Paradise strata originated near
North America (Tozer, 1982).

TRIASSIC COVER ON THE GOLCONDA ALLOCHTHON
[locs. 32, 41-43]
(Lander and Pershing counties, Nevada)

The Golconda Allochthon is a Carboniferous to mid-
Permian package of sedimentary and volcanic rocks
in thrust contact with an older, more eastern Paleozoic
allochthon (the Roberts Mountains Allochthon) or its
Upper Paleozoic cover. Triassic strata overlying the
Golconda Allochthon rest in unconformable contact
on a deformed Carboniferous to mid-Permian package
of deep-water pelagic rocks, turbidites, and pillow la-
vas (Silberling et a/., 1987) variously interpreted as a
collapsed back-arc basin or an accretionary prism (see
Miller et a/., 1984; and Brueckner and Snyder, 1985,
for discussions of each model).

Cyrtinoid spiriferinaceans occur within the Upper
Triassic Cane Spring Formation of the Star Peak Group
and the Osobb, Dun Glen, and (possibly) Winnemucca
formations of the Auld Lang Syne Group (as restricted
by Oldow, Bartel, and Gelber, 1990). The Cane Spring
includes carbonates that represent a range of shallow-
water environments (Nichols and Silberling, 1977),
whereas the overlying Auld Lang Syne clastic sedi-
ments were deposited in partly deltaic, to shallow sed-
imentary environments (Burke and Silberling, 1973).

Three lines of evidence suggest that the Golconda
Allochthon was emplaced against cratonal North
America during Late Permian to Early Triassic time.
First, stratigraphic evidence presented by Lupe and
Silberling (1985) suggests that cratonal, nonmarine
sediments of the Chinle Formation in the Great Basin
show sedimentary cycles correlative with those of the
marine Auld Lang Syne Group, which overlies the Gol-
conda Allochthon.

Second, the age of the youngest sediments in several
areas of the Golconda terrane has been used to con-
strain the timing of emplacement of the allochthon, an
event generally referred to as the Sonoma orogeny (Sil-
berling and Roberts, 1962; Snyder and Brueckner,
1983). In the Schoonover sequence, for instance, the
youngest basinal Paleozoic deposits are of Early Perm-
ian age, leading Miller et a/. (1984) to conclude that
thrust-faulting of the basin occurred in post-Early
Permian time. Burke and Silberling (1973) suggest that
emplacement occurred during the Late Permian or Ear-
ly Triassic, a conclusion that has been supported by
numerous other workers (e.g., Speed, 1977; Brueckner

and Snyder, 1985; Brueckner, Snyder, and Boudreau,
1987; Speed, Elison, and Heck, 1988). However, there
is considerable evidence that deformation of sediments
within the Golconda terrane may have been diachron-
ous (e.g., Brueckner and Snyder, 1985) and also that
the emplacement itself may have occurred at slightly
different times in various areas of Nevada. There is
also evidence for some post-Triassic movement of rocks
in both the Golconda and Roberts Mountains terranes
during later orogenic episodes (Ketner, 1984, 1990;
Ehman, 1985, p. 145; Little, 1987; Thorman et al.,
1990).

A third, and more compelling line of evidence for a
Triassic or earlier emplacement of the Golconda ter-
rane is that the Upper Triassic Auld Lang Syne clastics
were derived from an eastern, low-lying continental
source — possibly in the Colorado Plateau or Rocky
Mountain regions (Compton, 1960; Silberling and
Wallace, 1969; Burke and Silberling, 1973). The com-
position of these clastics differs from that expected for
western Nevada terranes derived from local sources
(Burke and Silberling, 1973). Thus, even if the timing
of emplacement was diachronous and spanned some-
what more than the Permian—Early Triassic range usu-
ally assigned to the Sonoma orogeny, an eastern source
for Upper Triassic Auld Lang Syne Group clastics
(Burke and Silberling, 1973) suggests that by Late Tri-
assic time the Golconda terrane adjoined the North
American craton (Lupe and Silberling, 1985).

EASTERN KLAMATH TERRANE
[locs. 1-5, 44]
(Shasta county, California)

The Eastern Klamath terrane, which represents a
Paleozoic—Early Mesozoic island-arc sequence in
northern California, preserves a Triassic through Mid-
dle Jurassic sedimentary section of carbonate, shale,
and coarse volcaniclastic sediments (Sanborn, 1960;
Silberling et a/., 1987). Cyrtinoid spiriferinacean bra-
chiopods occur in two formations within the Triassic
part of the section in Shasta County: the Hosselkus
Limestone and the Devil’s Canyon Member of the
Modin Formation. The Hosselkus consists of lime-
stones and shales and is probably in conformable con-
tact with the Pit Shale below, while it is only ques-
tionably conformable with the lithologically similar,
but more shaly, Brock Shale above. The Brock appears
to be in conformable contact with the Modin For-
mation above. Sanborn (1960) divided the Modin into
three subunits, from oldest to youngest, the Hawkins
Creek Member (volcanics and conglomerate), the De-
vil’s Canyon Member (tuffaceous limestone, lime-
stone, and calcareous sandstone), and the Kosk Mem-
ber (argillites, sandstones, and volcanics).

72 BULLETIN 337

IZEE TERRANE
[locs. 48, 49, 51]
(Suplee area, east-central Oregon)

Triassic strata in the Suplee area of east-central Or-
egon are assigned to the Begg and Brisbois formations
and the Rail Cabin Argillite (Dickinson and Vigrass,
1965). The Begg Formation, including mudstone, con-
glomerate, volcaniclastic rocks, lava, and rare bioclas-
tic limestone, rests unconformably on unnamed Pa-
leozoic rocks. The Begg contains possibly redeposited
mollusks and corals of probable early Karnian age and
has a conformable contact with the overlying Brisbois
Formation, in part of late Norian age. The Brisbois,
lithologically similar to the Begg, has produced all the
spondylospirid brachiopods recovered from the area.
It is unconformably overlain by the Rail Cabin Argil-
lite, of middle to late Norian age, which is overlain
concordantly by the Lower Jurassic Graylock For-
mation. Hettangian fossils in the basal strata of the
Graylock suggest that the uppermost beds of the Rail
Cabin could extend upward into the earliest Jurassic.
Silberling (written commun., 1990) suggests that ‘‘most
of the Late Triassic shelly fossils in the Begg and Bris-
bois are reworked,” and that “the Rail Cabin is evi-
dently a facies of part of the Brisbois, as interpreted
by Dave Taylor.”

WALLOWA TERRANE
flocs 125152212 28547]
(Wallowa County, Oregon, and Nez Perce and
Lewis counties, Idaho)

The Wallowa terrane, a Late Paleozoic—Early Me-
sozoic volcanic arc with overlying Triassic—Jurassic
limestones and clastic sediments, contains cyrtinoid
spiriferinacean brachiopods of both early and late No-
rian ages. Within this terrane, early Norian brachiopod
localities occur in Hells Canyon and in the Wallowa
Mountains, whereas the only late Norian locality is an
isolated limestone quarry, often called the Lewiston
locality (Stanley, 1979, p. 25), which is far to the north-
east of the other Triassic carbonate outcrops. The var-
ious Wallowa localities are discussed in further detail
below.

Early Norian (locs. 12, 21, 28, 47).—In the Wallowa
Mountains of northeastern Oregon, Triassic strata in-
clude the Martin Bridge and Hurwal formations, and

's informally designated the Lower Sedimentary Se-
[LSS] (Smith and Allen, 1941). The LSS in part
rlies, and in part may be equivalent to the largely

lover Creek Greenstone. The LSS is com-
conformably overlain by the Martin Bridge
Formation; and several hundred feet below that con-
tact the LSS contains fossils suggestive of the middle
upper Karnian Tropites welleri Zone. The Martin Bridge
Formation as the name is applied in Hells Canyon and

D mM +?
rermMian &

monly u

the northeastern Wallowa Mountains, consists largely
of shelf carbonates and includes typical early Norian
faunas. It contains the only spondylospirid brachio-
pods recovered from the area, and it is apparently con-
formably overlain by the Hurwal Formation. The Hur-
wal, in the northern Wallowa Mountains, contains
several ammonoid faunas of from early to middle No-
rian age (Silberling and Tozer, 1968), and it is overlain
unconformably by Tertiary volcanics (Reeside et ai.,
1957).

Late Norian (loc. 15).—An isolated limestone quarry
on the Nez Perce Reservation east of Lewiston, Idaho,
probably representing a roof pendant in the Wallowa
Batholith, contains scleractinian coral patch reefs with
an associated fauna of brachiopods, bivalves, and gas-
tropods. The coral-associated invertebrates represent
an epifauna-dominated assemblage; bivalves from this
locality consist of cemented and epibyssate (primarily
nestling) species (Newton, in prep.). The paleoecology
and structure of the scleractinian reefs has been doc-
umented by Stanley (1979).

The age of the carbonates at this locality is not well
documented. Although no ammonoids have yet been
recovered from this site, the reef-associated fauna seems
to indicate a late Norian age. Haas (1953) noted, in
particular, that the undescribed gastropod fauna at
Lewiston closely resembles late Norian gastropods from
Peru. Bivalves from the Lewiston site include no age-
diagnostic species. A diverse fauna of largely unde-
scribed brachiopods has been recovered from this site.
Two species of cyrtinoid spiriferinaceans are present,
one of which, S. /ewesensis (Lees, 1934), is of some
biostratigraphic utility. The other, so far as is known,
occurs only at this locality.

The spatial isolation of this limestone block makes
correlation with units in Hells Canyon (loc. 28) or the
Wallowa Mountains (locs. 12, 21, 47) difficult. Indeed,
if the late Norian age is correct, there are no time-
equivalent rocks of comparable lithology in Hells Can-
yon. Stanley (1979) has tentatively compared the Lew-
iston sequence with the late Norian Hurwal Formation,
though the lithofacies are not identical. Based on its
location, most workers have presumed that the Lew-
iston locality belongs to the Wallowa terrane, which
encompasses nearby parts of the Wallowa Mountains
and Hells Canyon areas (Silberling et a/., 1987).

STIKINE TERRANE
[locs. 29-31]
(Lake Laberge region, Yukon Territory, Canada)

The Stikine terrane is a Paleozoic through Early Me-
sozoic volcanic-are terrane characterized by volcanic
and sedimentary strata (Monger and Berg, 1987). The
Mesozoic rocks of the Lake Laberge region, Yukon
Territory, Canada, are divided into the Lewes River

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 73

Group (Triassic), the Laberge Group (Jurassic), and
the Tantalus Formation (Upper Jurassic or Lower Cre-
taceous). They are underlain by Permian limestones
of the Braeburn Formation. The Lewes River Group,
defined in this area by Lees (1934), has been divided
into seven formations, from oldest to youngest des-
ignated “A” through “G” (Tozer, 1958). These range
in age from Karnian through Norian. Formations “C”’,
“EE”, and ““G” consist mostly of limestone; ““B”’ and
“—D” are composed mostly of clastic rocks; and “A”
and ‘“‘F” contain both carbonate and clastic strata.
Spondylospirid brachiopods have been recovered only
from formations “‘F” and “G”’.

ALEXANDER TERRANE
[locs. 6, 7, 22-27]
(Gravina, Keku, Kuiu, and Screen Islands
and Cornwallis Peninsula, southeastern Alaska)

The Alexander terrane of southeastern Alaska rep-
resents a composite terrane with a complex Paleozoic
history that suggests that the three major subterranes
— Admiralty, Craig, and Annette — experienced dis-
parate tectonic regimes in pre-Late Paleozoic time
(Monger and Berg, 1987). Nonetheless, Pennsylvanian
plutons common to both the Alexander and Wrangellia
terranes indicate that at least part of the Alexander
terrane (and perhaps the entire composite terrane) was
united with the Wrangellia terrane at least by Late
Carboniferous time (Gardner et a/., 1988). A further
geographic constraint on the Alexander terrane comes
from paleomagnetic evidence that the terrane was lo-
cated far to the south in Pennsylvanian time (Van der
Voo et al., 1980), but had arrived at its present latitude
relative to North America by at least Triassic time
(Hillhouse and Grommé, 1980).

The Admiralty Subterrane, which contains the cyr-
tinoid spiriferinacean brachiopod localities herein dis-
cussed, includes Triassic strata of the Hyd Group and
the Keku Volcanics. The Hyd Group is divided into
the Burnt Island Conglomerate, Cornwallis Limestone,
Hamilton Island Limestone, and Hound Island Vol-
canics (Muffler, 1967). The Keku Volcanics consist of
felsic and mafic volcanic rock and intercalated clastic
units, which unconformably overlie the Permian Py-
bus Formation. The upper part of the Keku Volcanics
includes rare fossiliferous limestones, which contain
early to late Karnian faunas. The Burnt Island Con-
glomerate may be a lateral equivalent of the upper part
of the Keku Volcanics and at one locality contains [in
allochthonous pebbles] an early Karnian fauna. The
Cornwallis Limestone, of late Karnian to earliest No-
rian age, apparently overlies the Keku Volcanics con-
formably and the Burnt Island Conglomerate uncon-
formably. The Cornwallis is a gray oolitic limestone
containing faunas of late Karnian to earliest Nonan

age, and it is considered the shallow-water facies equiv-
alent of the deep-water Hamilton Island Limestone
(Muffler, 1967). Fossils from the Hamilton Island
Limestone suggest ages equivalent to that of the Corn-
wallis. Both the Cornwallis and Hamilton Island for-
mations are apparently conformably overlain by the
Hound Island Volcanics. This unit consists largely of
basaltic pillow breccia and pillow lava, andesitic vol-
canic breccia, and aquagene tuff, but includes some
thin-bedded limestone. Fossils from these limestones
suggest a late Karnian to late middle Norian age. The
Hyd Group is unconformably overlain by unnamed
Cretaceous lithic mudstones and sandstones. Silberling
(written commun., 1990) suggests that “the complex
facies relations implied by all of this probably indicate
some substantial tectonic disruption of these rocks.”
Cyrtinoid spiriferinacean brachiopods have been re-
covered from the Cornwallis Limestone and from the
Keku and Hound Island volcanics.

WRANGELLIA TERRANE
[locs. 13, 20]
(Wrangell Mountains, southern Alaska
and Nabesna quadrangle, south-central Alaska)

Triassic strata in the Wrangell Mountains of south-
central Alaska include the Nikolai Greenstone, the
Chitistone and Nizina limestones, and the McCarthy
Formation (Silberling and Tozer, 1968). Faunas from
shaly beds near the base of the Nicolai indicate a late
Ladinian to early Karnian age, while conformably
overlying the Nicolai, the Chitistone Limestone con-
tains 7ropites faunas near its base that are indicative
of an early late Karnian age. Fossils from beds tran-
sitional between the Chitistone and the overlying Ni-
zina Limestone suggest a latest Karnian to earliest No-
rian age. Fossils from the highest beds in the Nizina
suggest the Juvavites magnus Zone of late early Norian
age. Fossils in the lower part of the overlying McCarthy
Formation are typical of the Rhabdoceras suessi Zone
of late Norian age. Spondylospirid brachiopods have
been recovered in this area from beds transitional be-
tween the Chitistone and Nizina limestones and are
thus early Norian in age.

In the Nabesna quadrangle of south-central Alaska,
Triassic fossils are reported from three horizons. The
‘‘Upper Triassic limestone” of Richter (1976) contains
Monotis subcircularis Gabb, 1864, suggesting an early
late Norian age for that unit. Lying unconformably
below this unit, the ““Lower Triassic limestone” of
Richter has yielded spondylospirid brachiopods refer-
able to Spondylospira and has been assigned a late
Karnian age. This unit is bounded unconformably be-
low by the Nikolai Greenstone, an unfossiliferous unit
consisting of volcanics and volcanogenic sediments.

74 BULLETIN 337

The Nikolai is in turn bounded unconformably below
by the Eagle Creek Formation. The “Upper Triassic
limestone” is unconformably overlain by beds of Late
Jurassic to Cretaceous age which are assignable to the
“Nutzotin Mountains sequence” of Berg, Jones, and
Richter (1972).

Wrangellia is demonstrably an allochthonous Cor-
dilleran terrane, as attested by its distinctive internal
stratigraphy of Paleozoic arc rocks overlain by Triassic
basalt and Upper Triassic sedimentary strata (Jones,
Silberling, and Hillhouse, 1977; Jones et al., 1987),
and by its anomalous paleomagnetic signatures. Pa-
leomagnetic data suggest Triassic paleolatitudes of 14—
18°, whereas for the Alaska segment of Wrangellia the
expected Triassic paleolatitude would be approxi-
mately 35-37° (Hillhouse, 1977; Yole and Irving, 1980;
Hillhouse and Grommé, 1984; see compilations in
Stone and McWilliams, 1989). Recent paleomagnetic
modelling of the Jurassic and later motions of known
oceanic plates suggests that if Wrangellia accreted to
North America during Cretaceous time, Jurassic and
younger longitudinal displacement of Wrangellia with-
in the proto-Pacific ocean basin may have been as
much as 60° (Debiche, Cox, and Engebretson, 1987).

CHULITNA TERRANE
[loc. 53]
(Alaska Range, south-central Alaska)

The Chulitna terrane consists of Upper Devonian
ophiolite, overlain by Upper Paleozoic volcanic and
sedimentary rocks, Triassic carbonate, terrigenous
clastic, and basaltic rocks, and Jurassic clastic sedi-
mentary rocks and chert (Jones ef a/., 1980; Jones et
al., 1987). Within a complex Upper Triassic unit of
interstratified pillow basalt and limestone, Spondylo-
spira lewesensis (Lees, 1934) has been recovered in
association with large (up to 20 cm) megalodontid bi-
valves. Megalodontids are typical of shallow-water
platformal facies.

CERRO DE PASCO REGION
[loc. 16]
(Peru)

Spiriferina steinmanni Tilmann, 1917 and Spirifer-
ina sp. of Tilmann, 1917 (the latter here referred to
Spondylospira tricosta, n. sp.), are both referable to the

lylospirid genus Spondylospira Cooper, 1942 and
-ecovered from beds in the Triassic—Jurassic

section of Peru that Tilmann (1917, p. 644)

kes and Haas (1944, p. 284) referred to the
Szekeley and Grose (1972), in a detailed
e-examination of these beds at three localities in the
same region (Ninacaca; old trail connecting Carhua-
mayo and Pasco; Yanamarca), suggest that most of the
Jurassic Aramachay and Condorsinga formations are

liddie Lias.

missing here. Thus, the fossil-bearing sequence sam-
pled by Tilmann is probably best referred to the Cham-
bara Formation of the Pucara Group, which, on pa-
leontological grounds, is of Norian age. Lower Jurassic
(Sinemurian) ammonites recovered by Tilmann from
the nearby San Blas area were not found in association
with any spondylospirid brachiopods. At one set of
localities south of Cerro de Pasco, however, an un-
described species of Spondylospira is associated with
a sponge fauna that suggests a Jurassic age (Hoover,
1990). Thus, South America may be the only place
spondylospirids are not limited to the Upper Triassic.

SYSTEMATIC PALEONTOLOGY
INTRODUCTION

Abbreviations of Repository Institutions

The material discussed and described in this report
comes from several sources and collections. These bear
locality and/or catalogue numbers with acronyms as
follows:

CASG: California Academy of Sciences, Geology Col-
lections, San Francisco, CA.

GSC: National Type Collection of Fossil Invertebrates
and Plants, Geological Survey of Canada, Ottawa,
Ontario, Canada.

LSJU: Leland Stanford Junior University, Stanford,
CA.

USGS: United States Geological Survey, U.S. Na-
tional Museum of Natural History, Washington, DC.

USGS M: United States Geological Survey, Menlo
Park, CA.

USNM: U.S. National Museum of Natural History,
Smithsonian Institution, Washington, DC.

(other): Various collections, identified only by field
number, currently housed in the collections of the
U. S. Geological Survey, Denver, CO.

Measurement Abbreviations and Subscripts

In the tables of measurements: a.v. = articulated
valves; d.v. = dorsal valve; v.v. = ventral valve. Sev-
eral subscripts are used to qualify measurements. The
subscript . indicates that the measurement was esti-
mated and is probably correct only to the nearest mil-
limeter. The subscript . indicates that the specimen
was crushed in such a way as to render the measure-
ment non-representative; if . is also present, the value
given was estimated to correct this deficiency. The sub-
script ,, indicates that the specimen is broken and im-
plies that the missing portion is necessary to that mea-
surement; this may also be accompanied by the
subscript . with the same meaning as given above. A
measure across a structure with bilateral symmetry

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 75

(e.g., brachiopod hinge width) may be estimated by
doubling the half-measure [distance from symmetry
plane to distal extremity]. Use of this procedure is
indicated by the subscript ,, .

Terminology of Types

The terminology of types used in this report follows
that of Frizzell (1933). Of the types defined therein,
only the chorotype is likely to be unfamiliar to the
reader. Frizzell (p. 646) defined it as: “‘A fossil speci-
men collected from the same stratum as the type, but
from a neighboring locality.”

Size as a Taxonomic Character

Two samples of a continuous species population,
taken from temporally or spatially disjunct localities,
may exhibit different mean and range in individual
size. With such material, there is no way, a priori, to
determine whether one is dealing with two genetically-
distinct species or with a single size-variable one. In-
dividuals that differ on/y in size are here considered
conspecific; and that size as a basis for specific dis-
crimination, taken alone, is considered invalid. In many
monotypic new genera, however, the diagnosis of the
type species indicates the mean size of the individuals.
This implies that size is of significance in specific dis-
crimination.

Specific discrimination by size, however unrealistic,
is a necessary approximation (as are many paleonto-
logical species) in the initial stages of paleontological
investigation. The advantage of biological and pale-
ontological taxonomy is that it is not static, but is
cumulative, and may be emended as more data become
available. Thus, in this report, the type species of a
monotypic new genus may be diagnosed as “‘small’’,
based solely on the subjective opinion that other, larg-
er, possibly distinct congeneric species populations re-
main to be sampled and described.

““Lumpers” and “Splitters”

I do not think of myself as either a “splitter”, a
manufacturer of perhaps unnecessary new taxa, nor as
a “lumper’’, including new forms within the broad
potential morphological variation of a given taxon.
Several of the taxa reported herein are new genera
represented by single new species, and several of these
are represented by relatively few specimens from a
small number of localities. The relatively bizarre mor-
phological characters of many of these forms suggest
to me that they are distinct at the generic level, but
the thought remains that I do not have much material
on which to base this suggestion. I am left with the
task, which is the task of all taxonomists, of deter-
mining the closeness of relationship of the forms under
consideration to each other and to previously-de-

scribed ones, and have reluctantly decided that they
are indeed new, and not just on the fringes of variation
of existing forms. I am comforted by the belief that
taxonomic names, whatever else they are in genetic
terms, are “handles” for discussion of a definite group
of specimens, and that their correspondence to genetic
groups becomes increasingly vague as the temporal
distance from the Recent increases.

This would not be a problem if all I was doing was
describing morphology and making morphological
comparisons. I am, however, making paleobiogeo-
graphic inferences based on these taxonomic assign-
ments, and the taxonomic assignments favor the pa-
leogeographic arguments presented. I feel it necessary
to state that the taxonomic assignments presented
herein were made on morphologic grounds, with no a
priori thought to their paleobiogeographic conse-
quences, but also that I take full responsibility for those
assignments.

SYSTEMATICS
Order SPIRIFERIDA Waagen, 1883
Suborder SPIRIFERIDINA Waagen, 1883
Superfamily SPIRIFERINACEA Davidson, 1884
Family SPONDYLOSPIRIDAE, new family

Diagnosis.—Cyrtiniform spiriferinaceans with large
functional pedicle in the adult. Delthyrium open or
closed by deltidial structures. Additional posterior
covering plates (collectively the cooperculum; see Text-
fig. 2) rarely preserved. Dental plates converging an-
teriorly to join strong median septum as spondylium;
septum continuing posterior to junction of dental plates.
Jugal supports attached to valve floor. Spires of brach-
idium directed posterolaterally or ventrally.

Remarks.—The genera Spondylospira Cooper, 1942
and Vitimetula, Dagyspirifer, and Pseudospondylo-
spira, new genera, are included in the new family Spon-
dylospiridae. The erection of a new family is justified
by evidence that all four genera possessed a compar-
atively large functional pedicle in the adult stage, a
feature rare in other non-Paleozoic cyrtinoid spirifer-
inaceans. The size of the structure is indicated by the
area taken up by inferred pedicle foramena. A fora-
menate condition has not been reported in the related
family Laballidae, except for the two types of foramena
cited for the Thecocyrtellinae. Among the genera as-
signed to the Thecocyrtellinae (Thecocyrtella Bittner,
1892; Thecocyrtelloidea Yang and Xu, 1966; Klipstei-
nella Dagys, 1974), only Thecocyrtelloidea appears to
possess other than the normal single pedicle opening.
In Thecocyrtelloidea numerous “pedicle tubules” (Yang
and Xu, 1966, pl. 9, fig. 14, and pl. 10, fig. 2) appear
to pierce the deltidium throughout its length, a scheme
different from that seen in the Spondylospiridae.

76 BULLETIN 337

Views of the cooperculum of Spondylospira lewesensis (Lees, 1934). A, thin-section, x12, 11.7 mm dorsad of ventral beak,

sural plane, of ventral valve, GSC 69374D (chorotype, from loc. 30), showing ridged interarea, dental plates, and median

if rior portion of the spondylium, deltidial plates, and severely distorted cooperculum. Note that the shell structure of all

)f the shell is fibrous and laminar, while the cooperculum is clearly punctate externally and laminar internally, as is the

(shown only as fragment at extreme upper left of figure), except for the ridged portion of the interarea; B, posterior view, = 14.5,

tion of ventral valve, GSC 69373 (chorotype, from loc. 30), showing punctate surface of cooperculum, its irregular mesial suture (ticks
and bottom), and the lateral rugae and vertical furrows on its surface. (enlargement, upside down, of Pl. 9, fig. 21].

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 77

The proximal portions of the pedicle in the Spon-
dylospiridae are of two types, which form the basis for
definition of two new subfamilies, the Dagyspiriferinae
and the Spondylospirinae. Each subfamily is repre-
sented by two or more genera and species, which in
other respects exhibit considerable homeomorphy. In-
dividuals of genera currently assigned to the Laballidae
may, on closer examination, prove to be spondylo-
spirids rather than laballids.

Subfamily DAGYSPIRIFERINAE, new subfamily

Included genera.—Dagyspirifer, n. gen.; Pseudo-
sponadylospira, n. gen.

Diagnosis.—Spondylospirid spiriferinacean bra-
chiopods with paired pedicle foramena; foramena
ovate, internally rimmed by callus thickening. Shells
commonly globose, inflated, rarely pyramidal. Delti-
dial or additional posterior covering plates not ob-
served.

Remarks.—The Dagyspiriferinae may be distin-
guished from the Laballidae by their pedicle foramena,
from the Thecocyrtellinae by the number and size of
these pedicle openings, and the apparent lack of pos-
terior covering or deltidial plates in these forms, and
from the Spondylospirinae by the paired, rather than
multiple pedicle foramena, and the globose, rather than
pyramidal shell habit.

Text-figure 3.— Dorsoposterolateral view (prepared using a camera
lucida), x13, of ventral valve of Dagyspirifer fascicostata, n. gen.
and n. sp., USNM 450274 (paratype, from loc. 23), showing one of
the two small, paired, ovate pedicle foramena that pierce the apices
of both the ridged portion of the interarea and the dental plates. The
thin ridge that appears to bisect the foramen is the edge of a very
narrow deltidial plate. The dorsal edges of the spondylium are bro-
ken.

Genus DAGYSPIRIFER, new genus

Etymology of name.—Honoring A. S. Dagys, Lith-
uanian student of Triassic brachiopods.

Description.—Globose shells with subpentagonal
outline. Surface macro-ornament pauciplicate. Micro-
ornament of co-marginal growth lamellae and fine pus-
tules.

Ventral valve deep, swollen in lateral view, with
distinct sulcus bounded by largest costae. Sulcus pro-
duced anterodorsally as tongue producing W-shaped
commissure in anterior view. Beak produced consid-
erably posterior to hingeline, commonly incurved
strongly, reaching to commissural plane. Interarea low,
small, narrow, vertically ridged, bordered anteriorly
by row of hinge denticles. Delthyrium wide for the
subfamily, open, rimmed by narrow apparent deltidial
plates. Spondylium as for subfamily. Paired pedicle
foramena at delthyrial apex piercing apical portions of
both ridged interarea and dental plates (i.e., spondy-
lium) and opening into shell interior cavity on either
side of median septum (see Text-fig. 3).

Ventral interior surface radially plicate, reflecting ex-
terior ornament. Muscle impressions indistinct.

Dorsal valve unknown.

Type Species.— Dagyspirifer fascicostata, n. sp.

Diagnosis.—Globose Dagyspiriferinae with pauci-
costate radial macro-ornament, distinct ventral sulcus
floored by costae, and narrow ridged portion of inter-
area, and small, oval, paired pedicle foramena.

Occurrence.—As for species.

Comparison.— Dagyspirifer, n. gen. is superficially
most similar to Orientospira Dagys, 1965, from which
it differs in its costate, rather than smooth ventral sul-
cus, its delicate internal structures and its paired apical
pedicle foramena. It differs from other cyrtinoid spi-
riferinaceans in its markedly globose, rather than py-
ramidal form. It differs from the other known dagys-
piriferine, Pseudospondylospira, n. gen., in this respect,
in the much narrower ridged portion of its interarea,
and in its coarser macroornament.

Dagyspirifer fascicostata, new species
Plate 9, figures 1-9

new genus and new species of Early Norian brachiopod. Hoover,
1991, p. 394, fig. 2.

Etymology of name.—L. fascis = bundle + L. costata
= ribbed.

Description.—Medium-sized globose shells; outline
subpentagonal in dorsal view. Surface macro-orna-
ment of four to six coarse costae on each flank, ante-
riorly broadening from apex to commissure; low broad
smaller costae arising from flanks of larger costae,
broadening anteriorly. Micro-ornament of co-marginal
growth lamellae and fine pustules.

78 BULLETIN 337

Table 2.— Measurements (in mm) of type specimens of Dagyspirifer fascicostata, n. gen. and n. sp. See p. 74 for explanations of abbreviations

and subscripts.

hinge width maximum width length depth specimen type
locality 23
USNM 450274 (paratype) 93 18.4 14.7, 8.6 V.V.
USNM 450275 (paratype) 10.6 24.2 19.1,. 10.7 v.V.
USNM 450276 (holotype) 13.3 25.0 25.0 1322 V.V.

Ventral valve deep, swollen in lateral view; having
prominent flat-bottomed sulcus, floored by several very
weakly rounded broad costae, bordered by two most
prominent costae, these running distinctly from apex
to anterior commissure, there produced dorsally as
tongue. Commissure in anterior view W-shaped. Beak
produced considerably posterior to hingeline, com-
monly incurved to commissural plane. Interarea low,
small, about one-half maximum shell width, vertically
ridged, marginally rimmed by low, anteromesially-di-
rected hinge denticles. Delthryium wide, open, rimmed
by narrow apparent deltidial plates. Small spondylium
formed by ventral convergence of dental plates; plates
ridged on mesial surfaces, ridges reflecting previous
positions of anterior edges of dental plates during on-
togeny. Low median septum emergent from bottom

4.—Anterodorsolateral (interior) view (prepared using
x 13%, of ventral valve of Dagyspirifer fascicostata,
gen. and n. sp., USNM 450275 (paratype, from loc. 23), showing
» horizontal bar (at right) that connects the dental plates and lateral

vall. The opening above the bar connects to one of the two
I -d pedicle foramena that pierce both the ridged portion of the
interarea and the dental plate (see Text-fig. 3). The Y-shaped struc-
ture at lower left is the posterior portion of the spondylium.

mera 1ucica),

furrow of spondylium, produced a short distance an-
terodorsal of dental plates. Paired oval pedicle fora-
mena at apex of delthyrium opening into shell interior.

Ventral interior with thin median septum extending
about one-half valve length. Surface radially costate,
reflecting exterior macro-ornament. Thin, small, hor-
izontal plate joining apical dental plate to lateral shell
wall (Text-fig. 4). Muscle impressions indistinct.

Dorsal valve unknown.

For measurements, see Table 2.

Occurrence.—Dagyspirifer fascicostata, n. sp., has
been recovered only from locality 23 (Alaska, Corn-
wallis Limestone, Alexander terrane), from beds as-
signed to zone LNol of Tozer (1980), based on asso-
ciated fauna [Silberling, written commun., 1989].

Diagnosis.—Fascicostate Dagyspirifer.

Types.—Holotype, USNM 450276; paratypes,
USNM 4590274, USNM 450275.

Comparison.— Dagyspirifer is at present monotypic.
The fasciculate macro-ornament of D. fascicostata is
assumed to be of significance only at the species level.
D. fascicostata is probably more closely related to spe-
cies of Pseudospondylospira, n. gen. than to those of
Orientospira Dagys, 1965, even though it is superfi-
cially homeomorphic with the latter. It differs from
species of Orientospira in its paired pedicle foramena,
its more complex macro-ornament and its more del-
icate internal structures.

Genus PSEUDOSPONDYLOSPIRA, new genus

Etymology of name.—L. pseudo = false + Spon-
dylospira.

Description.—Subglobose, unequally biconvex, cos-
tate shells with modestly developed dorsal fold and
ventral sulcus. Ornament weaker on mesial than on
lateral shell surfaces. Outline subsemicircular in dorsal
view, with rounded dorsal umbo and pointed ventral
beak. Valves meeting at acute angles on flanks, at right
angles in mesial regions.

Ventral valve high. Costae broadening distally. In-
terarea duplex: laterally smooth, punctate, mesially
vertically ridged, impunctate; inner ridged portion lat-
erally rimmed by low serrate ridge in some specimens,
anteriorly bordered by hinge denticles that terminate
vertical ridges. Delthyrium high, narrow, open, with
narrow, marginal, disjunct apparent deltidial plates.
Paired ovate apical pedicle foramena piercing both

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 719

ridged interarea and apical portions of dental plates.
Low extensions of median septum protruding poste-
riorly from floor of spondylium.

Dorsal valve ornamented as for ventral. Umbo low,
broad.

Ventral interior surface reflecting exterior ornament.
Spondylial elements very thin, delicate. Paired ovate
apical pedicle foramena bearing raised callus rims.

Dorsal interior with bilobate anteroposteriorly stri-
ate cardinal process, flanked by anteriorly produced
articulatory shelf. Broad inner socket ridges flanking
wide, open notothyrium. Descending branches of spire
arising from dorsal margins of socket ridges, joined to
valve floor by narrow ridge. Smooth ovate paired ad-
ductor muscle scars. Surface reflecting external orna-
ment.

Type species.— Pseudospondylospira perplexa, ni. sp.

Diagnosis.—Subglobose Dagyspiriferinae with cos-
tate radial ornament, and large duplex interarea, and
relatively large, paired, oval apical pedicle foramena.

Occurrence.—As for species.

Comparison.—Pseudospondylospira, n. gen., out-
wardly resembles Spondylospira Cooper, 1942, very
closely. In specimens that preserve the apex of the
ventral valve, however, it may easily be distinguished
from Spondylospira, by its paired, ovate, rather than
multiple, slit-like pedicle foramena. It is otherwise dis-
tinguished from Spondylospira by its more delicate
shells, more globose, less pyramidal shape, and by the
limitation of the ridged portions of its interarea to a
smaller portion of the posterior face of the ventral
valve. Some specimens of two species of Spondylospira
[S. lewesensis (Lees, 1934) and S. parmata Hoover,
1983] preserve both deltidial plates and cooperculum,
and the shell structure of the ridged portion of the
interarea in Pseudospondylospira leads me to suggest
that a cooperculum was present in the latter genus as
well. Pseudospondylospira is distinguished from Da-
gyspirifer, n. gen., by its less globose form, its costate,
rather than paucicostate macro-ornament, its relative-
ly large, paired, oval pedicle foramena, and by its less
pronounced fold and sulcus development. Its differs
from laballid spiriferinaceans by having pedicle fora-
mena, and from thecocyrtelline spiriferinaceans by
these foramena being paired, rather than single or mul-
tiple, and by having an open delthyrium.

Pseudospondylospira perplexa, new species
Plate 10, figures 14-24

new genus and new species of Late Karnian to Late Norian bra-
chiopod. Hoover, 1991, p. 394, fig. 3.

Etymology of name.—L. perplexus = obscure, am-
biguous.

Description.—Subglobose, inequibiconvex, costate
shells with modest development of dorsal fold and

ventral sulcus. Costae weaker, narrower, more closely
spaced on fold and sulcus than on flanks. Shell outline
subsemicircular in dorsal view, with rounded dorsal
umbo and pointed ventral beak. Commissural angle
acute on flanks, about 90° mesially.

Ventral valve transverse, wider than long; high,
rounded, with tongue-like semicircular dorsal exten-
sion of mesial sulcus. Costae low, rounded, gradually
broadening distally. Small, pointed beak hooked pos-
terior to hingeline. Interarea duplex; laterally smooth,
mesially vertically ridged; in some specimens rimmed
by low serrate ridge on lateral margins of mesial ridged
portion; anteriorly bordered by articulatory hinge den-
ticles. Delthyrium high, narrow, open, with narrow,
marginal, disjunct apparent deltidial plates. Paired
elongate oval pedicle foramena piercing ridged inter-
area and apical portions of dental plates. Spondylium
floored mesially by thin posterior extension of median
septum; septum projecting somewhat dorsad of edges
of dental plates.

Dorsal valve bearing moderately strong to weak
macro-ornament of radial costae; costae more narrow,
weak, and closely spaced on fold than on flanks, in-
creasing in number anteriorly by bifurcation, median
costa rarely broader or higher than adjacent costae.
Anterior commissure in dorsal view mesially resected
by sulcal tongue of opposite valve. Umbo low, broad,
produced slightly posterior to hinge; incurved only
slightly ventrad to hingeline.

Ventral interior surface reflecting exterior ornament
of costae, smooth distal interarea and ridged mesial
interarea. Elements of spondylium (dental plates, me-
dian septum) thin, delicate. Pedicle foramena bearing
raised callus rims; muscle impressions otherwise in-
distinct.

Dorsal interior with bilobate, anteroposteriorly stri-
ate cardinal process, flanked by anteriorly-produced
articulatory shelf. Shelf bearing denticulate dorsad step
at about mid-length; narrow lengthwise furrows pos-
terior to step; deep, oval articulatory fossae just an-
terior to step. Broad inner socket ridges flanking wide,
open notothyrium. Descending branches of spire aris-
ing from dorsal margins of socket ridges, joined to
valve floor in narrow ridge at about one-half valve
length. Median myophragm separating smooth, ovate,
paired adductor scars; myophragm variably devel-
oped, low; may extend anteriorly about one-fifth valve
length. Interior surface radially striate, reflecting ex-
terior ornament.

For measurements, see Table 3.

Occurrence.— Pseudospondylospira perplexa, Nn. sp.,
has been recovered in the present collections from lo-
calities 7 [Alaska, Cornwallis Limestone, Alexander
terrane, zones UCI-LNol(?) of Tozer (1980), based
on associated fauna], 18 [Nevada, Jungo terrane, zone

80 BULLETIN 337

Table 3.—Measurements (in mm) of type specimens of Pseudospondylospira perplexa, n. gen. and n. sp. See p. 74 for explanations of

abbreviations and subscripts.

eS owoo.._“

striate hinge maximum
width interarea width width length thickness specimen type
ee ee SSS SS SSS Ss
locality 22
USNM 450277 (paratype) — Ips 20 24 15:3 a.v.
USNM 450278 (holotype) 11.0 15.6 28, 24 22.5 a.v.

LNo3 of Tozer (1980), based on associated fauna], 22
[Alaska, Cornwallis Limestone, Alexander terrane,
zones UCI-—UNol of Tozer (1980), based on associ-
ated fauna], 23 [Alaska, Cornwallis Limestone, Alex-
ander terrane, zone LNol of Tozer (1980), based on
associated fauna], 24 [Alaska, Cornwallis Limestone,
Alexander terrane, zones UC1-—LNo2 of Tozer (1980),
based on stratigraphic position], 25 [Alaska, limestone
bed in Hound Island Volcanics, Alexander terrane,
zones UC2-LNol of Tozer (1980), based on associated
fauna], 27 [Alaska, Cornwallis Limestone, Alexander
terrane, zones UCI-UNol of Tozer (1980), based on
stratigraphic position], 28 [Oregon, Martin Bridge For-
mation, Wallowa terrane, zone LNol of Tozer (1980),
based on associated fauna], and 41 [Nevada, Osobb
Formation, Triassic cover on the Golconda Alloch-
thon, zones LNo2—LNo3 of Tozer (1980), based on
stratigraphic position]. See Table 1 for Tozer’s (1980)
zonal scheme.

Diagnosis.—Transverse Pseudospondylospira with
low dorsal fold and shallow ventral sulcus.

Types.—Holotype, USNM 450278; paratypes,
USNM 450277, USNM 450279, USNM 450280,
USNM 450281, USNM 450282.

Comparison.— Pseudospondylospira perplexa, n. sp.,
is at present the only species in the genus. The as-
sumption is made that the outline and strength of fold
and sulcus development are specific rather than generic
characters. Mean size may also prove to be significant,
but no conclusions can be made in the absence of large
population samples of P. perplexa or specimens of oth-
er congeneric species. Several specimens from a single
locality in Oregon have been tentatively assigned to
this species (see below). Their stronger macro-orna-
ment and more pronounced fold and sulcus develop-

“nt are near the limit of what I would consider nor-

pecific variation.
Pseudospondylospira perplexa, n. sp.,
ified from silicified material, which com-
nly preserves the subfamilially diagnostic apex of
ventral valve. Calcareous “‘crackout’”’ specimens,
however, commonly lack this portion of the shell, and
levelopment of such specimens from matrix involves
ngthy and complex preparation. I have not identified
specimens as P. perplexa unless, in addition to other
criteria, the diforamenate condition could be ascer-

tained. Hence, many individuals that may be P. per-
plexa have been assigned to less restrictive taxonomic
categories, and the cited occurrence of the species may
not be representative of its true geographic and strati-
graphic range.

Pseudospondylospira aff. P. perplexa, new species
Plate 10, figures 25-27

Remarks.—A small partial ventral valve and a larger
complete dorsal valve of a form closely related to P.
perplexa were recovered from locality 28 (Oregon,
Martin Bridge Formation, Wallowa terrane) in beds
assigned, on the basis of associated fauna, to zone LNol
of Tozer (1980) by Silberling (written commun., 1989).
The paired pedicle foramena clearly place these spec-
imens within the Dagyspiriferinae, and the subglobose
form and comparatively subdued macro-ornament
suggest Pseudospondylospira. The macro-ornament is,
however, stronger than that in P. perplexa, and is ac-
companied by more pronounced development of the
fold and sulcus. The range of variation in macro-or-
nament observed in P. perplexa (see Pl. 10, figs. 20,
21) suggests this form may also lie within P. perplexa,
but too few specimens are present to permit more con-
clusive identification than the above.

Types.— Figured specimens, USNM 450283, USNM
450284.

Subfamily SPONDYLOSPIRINAE, new subfamily

Included genera.—Spondylospira Cooper, 1942; Vi-
timetula, n. gen.

Diagnosis.—Spondylospirid spiriferinacean bra-
chiopods with more than two pedicle openings. Del-
tidial plates and coopercula may be present. Shells
commonly subpyramidal, apically hooked, rarely glo-
bose or inflated.

Remarks.—The Spondylospirinae may be distin-
guished from the Laballidae by their pedicle opening,
from the Thecocyrtellinae by the size of that opening,
from the Dagyspiriferinae by their multiforamenate
rather than diforamenate pedicle opening, and in some
species of Spondylospira from all others by the pres-
ervation of two distinct kinds of coopercula (see Hoo-
ver, 1983; and Text-fig. 2 and Plate 10, figs. 10-13
herein).

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 81

Genus SPONDYLOSPIRA Cooper, 1942

Type species.— Spondylospira reesidei Cooper, 1942,
p. 232; Cooper, in Shimer and Shrock, 1944, p. 359,
pl. 140, figs. 43-47.

Diagnosis.—Spondylospirine spondylospirids with
weak to strong dorsal fold and ventral sulcus, bearing
ornament of strong costae on flanks and weaker costae
on fold and sulcus. Interarea vertically ridged; del-
thyrium commonly preserved as open, in some spec-
imens covered by deltidial plates or deltidium; inter-
area in some specimens shielded by multipartite shield
(cooperculum). Hinge denticulate. Ventral interior with
spondylium basally bisected by median septum; dorsal
interior with descending branches of spire jugate an-
teriorly, supported dorsally by meshwork connected to
valve floor.

Occurrence.—Spondylospira occurs in beds of from
Karnian to Norian age in North and South America.
Spondylospira cf. S. lewesensis (Lees, 1934) is the only
species referred to the genus reported from outside the
Western Hemisphere [by Dagys (written commun.,
1990) from the Koryak mountains of the northeast-
ernmost U.S.S.R.]. This species was originally re-
ferred to Spondylospira alia (Hall and Whitfield, 1877)
by Bychkov and Dagys (1984, p. 13).

Comparison.—Well-preserved specimens of Spon-
dylospira may be distinguished from other non-spon-
dylospirine cyrtinoid spiriferinaceans by their multiple
apical pedicle foramena. Less well-preserved speci-
mens may be distinguished from Zugmayerella Dagys,
1963, Laballa Moisseiev in Dagys, 1962, Pseudola-
balla Dagys, 1974, Spinolepismatina Dagys, 1974, and
Orientospira Dagys, 1965, by their costate fold and
sulcus. The latter characters also distinguish Spondy-
lospira from the other known spondylospirine, Viti-
metula, n. gen.

Discussion.— Vokes and Haas (1944) referred the
following species to Spondylospira: Cyrtina lewesensis
Lees, 1934, Spiriferina acrotamboensis Korner, 1937,
Spirifera (Spiriferina) alia Hall and Whitfield, 1877,
Spiriferina canavarica Tommasi var. robusta Korner,
1937, Spiriferina koessenensis Zugmayer, 1882, Spi-
riferina steinmanni Tilmann, 1917, Spiriferina sp. of
Tilmann, 1917, Spiriferina(?) sp. indet. of K6rner, 1937,
and Spondylospira reesidei Cooper, 1942.

Cyrtina lewesensis Lees, 1934 is the name first ap-
plied to the form which typifies Spondylospira. Widely
variable in size, strength of costation, and number of
costae, this species is the senior synonym of both Spi-
riferina acrotamboensis [Pl. 9, figs. 27-29], and Spon-
dylospira reesidei [Pl. 9, figs. 22-26]. Hoover (1983)
reported that preparation of the ventral valve which
is the holotype of S. (S.) alia Hall and Whitfield, 1877
[USNM 12671; Pl. 10, figs. 4-7] reveals a shell that
may not even be punctate, a low, non-ridged interarea,

non-denticulate hinge, and absence of the “‘cyrtinoid”’
spondylium that is the uniting characteristic of the
laballids and spondylospirids. S. (S.) alia cannot be
placed within either family, and only questionably fits
in the Spiriferinacea. The specimen figured by Cooper
(in Shimer and Shrock, 1944, pl. 141, figs. 48-51) as
Spondylospira alia (Hall and Whitfield, 1877) is here
placed in synonymy with S. /ewesensis (Lees, 1934).
Specimens of S. canavarica robusta, S. steinmanni,
and S. (?) sp. indet. of K6rner, were not available for
comparison, but from the published illustrations sug-
gest assignment to Spondylospira. Spiriferina sp. of
Tilmann (1917) is herein referred to Spondylospira tri-
costa, n. sp. Spiriferina koessenensis was removed from
Spondylospira by Dagys (1963), who defined it as the
type species of Zugmayerella.

Spondylospira lewesensis (Lees, 1934)
Plate 9, figures 10-38; Plate 10, figures 1-3

Cyrtina lewesensis Lees, 1934, p. 35, pl. 1, figs. 14-16.

Spiriferina acrotamboensis Korner, 1937, p. 168, pl. 11, figs. 5-8.

Spondylospira reesidei Cooper, 1942, p. 232; Cooper, in Shimer and
Shrock, 1944 [part], p. 359, pl. 140, figs. 43-47 [USNM 103468a-
c, e-g only; not USNM 103468d (= Vitimetula parva, n. gen. and
n. sp.)]; Dagys, 1974, pl. 54, figs. 5-8; Hoover, 1983, figs. 3G-3K.

Spondylospira alia (Hall and Whitfield). Cooper, in Shimer and
Shrock, 1944, p. 359, pl. 140, figs. 48-51 [not Spirifera (Spiriferina)
alia Hall and Whitfield, 1877, p. 281, pl. 6, fig. 17].

Spondylospira aff. S. alia Hall and Whitfield [sic]. Rangel Z., 1978,
p. 25, pl. 1, figs. 3-5 [not Spirifera (Spiriferina) alia Hall and
Whitfield, 1877, p. 281, pl. 6, fig. 17].

Spondylospira lewesensis Lees. Tozer, 1962, p. 27, pl. 12, figs. 11-
13; Dagys, 1974, pl. 51, figs. 9a—b; Hoover, 1983, pp. 1026-1028,
figs. 3L-3S, 4E, 4F, 5.

Spondylospira aff. S. acrotamboensis Korner [sic]. Rangel Z., 1978,
p. 25, pl. 1, fig. 6, pl. 2, figs. 1-4.

Spondylospira cf. S. acrotamboensis Korner [sic]. Rangel Z., 1978,
p. 24, pl. 1, figs. 1, 2.

Description.—Spondylospira lewesensis (Lees) has
been amply described in the above accounts. Its dif-
ferences from related species are detailed under Com-
parison below. For measurements, see Table 4.

Occurrence.—Spondylospira lewesensis (Lees) has
been recovered from deposits of from early Karnian
to late Norian age. In the present collections, it is rec-
ognized from localities 5 [California, 7Modin Forma-
tion, Eastern Klamath terrane, zones LNol1—UNo3 of
Tozer (1980), based on associated fauna], 6 [Alaska,
Alexander terrane, zones UC1—UC3 of Tozer (1980),
based on associated fauna], 10 [Nevada, middle mem-
ber of Gabbs Formation, Paradise terrane, zones
UNo2-UNo3 of Tozer (1980), based on associated
fauna], 13 (Alaska, Wrangellia terrane, age unknown),
15 [Idaho, Wallowa terrane, zones (?) UNol-—UNo3 of
Tozer (1980), based on associated fauna (see Appendix
for discussion of age uncertainty at this locality)], 16
[Peru, zones UCI-—UNo3 of Tozer (1980), based on
associated fauna], 21 [Oregon, Martin Bridge Forma-

82 BULLETIN 337

Table 4.—Measurements (in mm) of specimens of Spondylospira lewesensis (Lees, 1934). See p. 74 for explanations of abbreviations and

subscripts.
dorsal costae on
hinge width maximum width length thickness fold/flank specimen type
locality 15
USNM 103468a' 9.1 13.7 8.4. 9.0. — v.v.
USNM 103468b? 14.0, 15.0, 9.0, 3.0. 6/6 d.v.
USNM 103468c? 14.1 14.1 6.1 3.0 4/6 d.v.
USNM 103468d? 10.1 12.4 8.8 6. — V.V.
locality 29
GSC 9619 (lectotype) 17.9 22.6 18.5 20.7 4/6 a.v.
GSC 9619a (paralectotype) 17.0 2257 20.7 21 6/6 a.v.
locality 30
GSC 69373 (chorotype) 14.9 22.4 23.8 17.8 6/6 a.v.
locality 39
CASG 66253.02 6.8 7.9 6.4 6.3 4/6 a.v.
CASG 66253.03 8.3 11.0 8.5 9.0. 4/6 a.v.
CASG 66253.04 10.4, 12.3 9.1 9.2 4/6 a.v.
CASG 66253.01 1S 14.0 10.0 9.5 6/7 a.v.
locality 45
CASG 66258.05 6.8 8.3 etl 5.8 4/4 a.v.
CASG 66258.01 7.6 8.6 5.4 6.2 4/4 a.v.
CASG 66258.06 6.8 8.4 5.6 5.0 4/4 a.v.
CASG 66258.07 8.0 9.2 7.5 6.3 4/7 a.v.
CASG 66258.08 7.9 9.7 6.8 8.3 = a.v.
CASG 66258.03 8.6 10.2 8.0 ed 4/6 a.v.
CASG 66258.02 10.4 10.5 6.5 6.9 4/6 a.v.
CASG 66258.09 10.4 11.5 8.2 8.0 4/6 a.v.
CASG 66258.10 10.8 12.8 8.9 8.0 4/8 a.v.
CASG 66258.04 13.2 13.8 8.5 9.8 6/6 a.v.

' Holotype of Spondylospira reesidei Cooper, 1942.
? Paratype of Spondylospira reesidei Cooper, 1942.

tion, Wallowa terrane, zone UC3 of Tozer (1980), based
on associated fauna], 26 [Alaska, limestone bed in Keku
Volcanics, Alexander terrane, zones LCI-UNo?2 of
Tozer (1980), based on associated fauna], 27 [Alaska,
Cornwallis Limestone, Alexander terrane, zones UC 1—-
UNol of Tozer (1980), based on stratigraphic posi-
tion], 29 [Yukon, Formation “‘F” of Lewes River
Group, Stikine terrane, zone UNo2 of Tozer (1980),
based on associated fauna], 30 [Yukon, Formation “F”
of Lewes River Group, Stikine terrane, zone UNo2 of
Tozer (1980), based on associated fauna], 31 [Yukon,
Formation “F” of Lewes River Group, Stikine terrane,
‘one UNo2 of Tozer (1980), based on associated fau-
3 | Nevada, lower member of Luning Formation,
terrane, zone LNol of Tozer (1980), based

it this stratigraphic level], 35 [Nevada, Lun-

ag Formation, Paradise terrane, zone LNo3 of Tozer
980), based on associated fauna], 36 [Nevada, Lun-
ing Formation, Paradise terrane, zone LNol1 of Tozer
(1980), based on fauna at this stratigraphic level], 37
[Nevada, lower Luning Formation, Paradise terrane,
zones LNol—LNo2 of Tozer (1980), based on fauna at
this stratigraphic level], 39 [Nevada, Jungo terrane,

zone LNo3 of Tozer (1980), based on associated fau-
na], 40 [Nevada, Luning Formation, Paradise terrane,
zone LNo3 of Tozer (1980), based on stratigraphic
position], 41 [Nevada, Osobb Formation, Triassic cov-
er on the Goldconda allochthon, zones LNo2—LNo3
of Tozer (1980), based on stratigraphic position], 45
[Nevada, Jungo terrane, zone MNo2 of Tozer (1980),
based on associated fauna], 47 [Oregon, Martin Bridge
Formation, Wallowa terrane, zones LNol—-MNo2 of
Tozer (1980), based on associated fauna], and 53 [Alas-
ka, Chulitna terrane, zone UC3 of Tozer (1980), based
on associated fauna]. This indicates a range of zones
LC1 through UNo3 of Tozer (1980), with ages in the
older end of the range most common in the Alexander
terrane. The early and middle Karnian extension of
the range is supported by only a single locality (loc.
26), and Dagys (written commun., 1990) doubts that
the species really ranges down into the Karnian at all.
Occurrences that may be Karnian are limited to the
Alexander and Chulitna terranes of Alaska.
Diagnosis.—Spondylospira with strong dorsal fold
costae that increase initially by equal bifurcation, sub-
sequently by equal or unequal bifurcation; widest at or

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 83

slightly anterior to hingeline; fold and sulcus moder-
ately to strongly developed; bears posterior cooper-
culum on interarea, but cooperculum rarely preserved.

Types.—Lectotype, GSC 9619; paralectotype, GSC
9619a; chorotype, GSC 69373; chorotype of S. acro-
tamboensis Korner, 1937, USNM 450285; figured
specimens, CASG 66248.01, CASG 66248.02, CASG
66251.01, CASG 66253.01, CASG 66258.01—CASG
66258.04, USNM 103468a, USNM 103468b, USNM
103468f, USNM 106521, USNM 450286, USNM
450287; measured specimens, CASG 66253.01-—CASG
66253.04, CASG 66258.01-CASG 66258.10, GSC
9619, GSC 9619a, GSC 69373, USNM_ 103468a-
USNM 103468d.

Comparison.— Spondylospira lewesensis (Lees, 1934)
includes forms that vary considerably in most of the
characters which define the species. It may most easily
be distinguished from S. tricosta, n. sp., by the manner
of initial increase in number of costae on the dorsal
fold. S. tricosta adds a single subordinate costa on ei-
ther side of a single initial costa, so that bilateral tri-
furcation takes place. S. /ewesensis simply bifurcates
the initial costa, and at least in the early ontogenetic
stages, bears an even number of dorsal fold costae, as
new ones are added symmetrically. S. parmata Hoo-
ver, 1983 may fall within the range of S. /ewesensis. It
was identified as a distinct species primarily because
(1) it is close to the limits of variation of S. /ewesensis
and (2) it exhibits features of particular interest that
are not well shown in S. /ewesensis, although they may
be present in some individuals of that species. S. par-
mata may be distinguished by its weak development
of the median fold and sulcus, and by its very numer-
ous, thin, closely spaced costae. It bears a cooperculum
that appears to be tripartite, as opposed to the bipartite
set of elements seen in some specimens of S. /ewesensis.
S. lewesensis includes multicostate forms referred by
Cooper to S. alia (Hall and Whitfield, 1877). Hoover
(1983) recognized that the type of Hall and Whitfield’s
species was not a spondylospirid.

In Spondylospira the dorsal interior spire support
meshwork memorialized by the generic name is not
always well-preserved. The beautiful silicification seen
in the types of S. acrotamboensis (KOrner, 1937) and
S. reesidei Cooper, 1942 is the exception rather than
the rule. Thus specific characters have tended to be
external. The types of S. reesidei (USNM 103468a-c,
e-g) however, suggest how variable such external char-
acters may be. In addition, it seems subjectively rea-
sonable that individuals of this elusive genus from such
geographically widely-separated regions as Peru, Ida-
ho, and the Yukon Territory of Canada might, in the
absence of other collections, belong to distinct species.
Thus, S. /ewesensis (Lees) from the Yukon Territory
of Canada was considered distinct from the Peruvian

species S. acrotamboensis (K6rner, 1937), and both of
those were considered distinct from S. reesidei Cooper.
These three species can only be differentiated as fol-
lows: S. acrotamboensis and S. lewesensis are larger,
and have more lateral costae, than does S. reesidei.
This distinction, as suggested above, is insufficient ba-
sis for specific discrimination when a large database is
available. S. /ewesensis and S. acrotamboensis have
more (4—6 vs. 4—5) lateral costae than does S. reesidei.
The latter phenomenon can be related directly to size.
Discounting these differences, the three species above
must be considered to form a single, externally vari-
able, hemispherically cosmopolitan species, Spondy-
lospira lewesensis (Lees).

Discussion.—The type material of S. reesidei in-
cludes individuals with two distinct types of ornament.
Most bear strong, commonly sinuous costae, that bi-
furcate anteriorly. A single ventral valve [USNM
103468d] bears only simple, very weak costae, and is
comparatively smooth. Examination of topotypic ma-
terial [loc. 15] has produced further specimens of this
smooth form. As no transitional forms occur, the
smooth form is here assigned to a distinct taxon. As
it lacks a costate fold or sulcus, it cannot be assigned
to Spondylospira [sensu stricto], and is therefore here
placed in the new genus Vitimetula. Ventral valves of
both forms show the multiple apical foramena char-
acteristic of the new subfamily Spondylospirinae.

Spondylospira parmata Hoover, 1983
Plate 10, figures 8-13

Spiriferina cf. S. suessi Winkler. Sanborn, 1960, pp. 20-21, pl. 2,
figs. 7, 8 [not Spiriferina suessi Winkler, 1859 = Laballa Moisseiev
in Dagys, 1962].

Spondylospira parmata Hoover, 1983, pp. 1025, 1026, text-figs. 3A—
3F; Hoover, 1991, p. 395, fig. 4.

Description.— Pyramidal, inequibiconvex costate
shell, with weak dorsal fold and ventral sulcus. Costae
increasing in number distally by bifurcation. Interarea
about half-covered by multipartite cooperculum, ex-
posed apically and basally.

Ventral valve about equal in length, width, and depth.
Outline triangular in lateral, anterior, posterior, or ven-
tral aspects. Surface macro-ornament of closely spaced
fine costae; at 10 mm from beak, about 15 on each
flank, and six narrower ones in shallow sulcus. Inter-
area mostly planar, but beak strongly hooked poste-
riorly at apex. Interarea surface vertically finely ridged;
about 17 ridges in a 5 mm distance at base. Hingeline
denticulate; denticles serving articulatory function.
Delthyrium high, open, narrow.

Cooperculum of single basal and two lateral plates,
together covering entire width of basal two-thirds of
interarea, exposing only oval apical mesial and low,
transverse slitlike basal areas. Surface rugose, with ap-

84 BULLETIN 337

Table 5.—Measurements (in mm) of the holotype of Spondylospira parmata Hoover, 1983. See p. 74 for explanations of abbreviations and

subscripts.

maximum
hinge width width

interarea height length thickness specimen type

locality 44

CASG 60975.01 (holotype) 21 be 2 ye

parent suture-lines demarcating basal one or two plates,
and disjunct right and left lateral plates. Basal plate
relatively smooth, lateral plates co-marginally rugose
around pedicle opening, bearing coarse rounded ver-
tical grooves on lateral plates. Lateral plates thickest
(ca. 2 mm) around pedicle opening, thinning distally
and toward the hingeline.

Dorsal valve with costate ornament in number and
character like that of opposite valve. Low mesial fold
developed. Hinge area bearing longitudinally-ridged
articulatory shelf and striate cardinal process.

Ventral interior with spondylium of dental plates,
supported and divided longitudinally by median sep-
tum. Interiors of either valve not otherwise observed.

For measurements, see Table 5.

Occurrence.—Only a single specimen confidently
identifiable as S. parmata Hoover, 1983, is known. It
was recovered from locality 44 (California, Devil’s
Canyon Member of the Modin Formation, Eastern
Klamath terrane), in beds assigned to zones UNol-
UNo3 of Tozer (1980) on the basis of associated fauna
(Silberling, written commun., 1989). Several other
specimens which appear conspecific but have not pre-
served the cooperculum characteristic of S. parmata
have been recovered from locality 44, and from locality
2 (California, 7Modin Formation, Eastern Klamath
terrane). Beds at the latter locality may be assigned to
zones MNol through UNo2 of Tozer (1980) based on
associated fauna (Silberling, written commun., 1989).

Diagnosis.—Equidimensional Spondylospira with
numerous fine costae, weak fold/sulcus development,
and a tri-partite cooperculum covering portions of the
ventral interarea.

Types.— Holotype: CASG 60975.01.

Comparison.—S. parmata may fall within the ex-
tremely broad range of variation of S. /ewesensis (Lees),
as discussed above. It is, however, near the limits of

yee of that variation, and its numerous fine costae

fold and sulcus development distinguish it

; of S. acrotamboensis, S. lewesensis, and S.

{t is more similar in exterior ornament to
spondylospira perplexa, n. sp., but differs from

form in having an interarea that covers the max-
num width of the shell, and in having multiple rather
han paired pedicle foramena. Its tripartite coopercu-
jum distinguishes it from all previously described spe-
cies of the genus. Topotypic specimens of S. /ewesensis
(Lees) also bear a cooperculum, which in that species

20.3 23. 29.2 ay.

appears to be comprised of only two paired lateral
plates.

Spondylospira tricosta, new species
Plate 11, figures 1-15

Spiriferina sp. Tilmann, 1917, p. 689, pl. 25, figs. 12a, 12b.

Etymology of name.—L. tri = three + L. costa =
rib.

Description.—Small, inequibiconvex, paucicostate
shells with distinct, costate dorsal fold and ventral sul-
cus; commonly high, thicker than long; transverse,
wider than long.

Ventral valve outline subsemicircular in ventral as-
pect, dorsally tongued in anterior aspect; pointed beak
commonly projecting posteriorly. Distinct median sul-
cus commonly bearing two weak radial costae; flanks
bearing four to five stronger costae; mesial costae only
originating at beak; others originating further dorsad
on flanks of posterior margin. Interarea planar to con-
cave posteriorly, bounded laterally by raised rim; ver-
tically finely ribbed, ribs terminating dorsally in fine
hinge denticles. Delthyrium narrow, open. Micro-or-
nament of fine co-marginal growth lines and radial
striae.

Dorsal valve anteriorly truncate in lateral aspect;
hinge width commonly slightly less than maximum
width, subequal to length. Broad median fold com-
monly bearing three costae on crest; mesial costa
strongest, all arising about four mm anterior to beak
by trifurcation. Flanks commonly bearing fine costae.

Ventral interior with spondylium and posteriorly-
produced median septum. Hinge teeth low, feeble, ar-
ticulation chiefly by hinge denticles.

Dorsal interior unknown.

For measurements, see Table 6.

Occurrence.—Spondylospira tricosta, n. sp., is known
only from the early Norian [zones LNol through LNo3
of Tozer (1980)]. It has been recognized in the present
study at localities 8 [Nevada, lower member of Luning
Formation, Paradise terrane, zone LNol of Tozer
(1980), based on associated fauna], 28 [Oregon, Martin
Bridge Formation, Wallowa terrane, zone LNol of
Tozer (1980), based on associated fauna], 32 [Nevada,
Osobb Formation, Triassic cover on the Golconda AI-
lochthon, zones LNol1-LNo3 of Tozer (1980), based
on stratigraphic position], 40 [Nevada, Luning For-
mation, Paradise terrane, zone LNo3 of Tozer (1980),
based on stratigraphic position], 41 [Nevada, Osobb

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 85

Table 6.— Measurements (in mm) of specimens of Spondylospira tricosta, n. sp. See p. 74 for explanations of abbreviations and subscripts.

costae on dorsal

hinge width maximum width length thickness fold/flank specimen type

locality 8

USNM 450288 17.0 2273. 17.6, Zee 5/5 a.v.
locality 32

CASG 66247.01 (holotype) Wey 13.8 11.4 11.5 3/5 a.v.

CASG 66247.02 (paratype) 10.6 15.4 11.5, 12.6 3/5 a.v.

CASG 66247.03 (paratype) 10.2 17.3 13:0; 13.8 3/5 a.v.

CASG 66247.04 (paratype) 14, 18.5 13.9 15.8 4/5 a.v.
locality 40

CASG 66254.01 11.1 14.8 12.0 15.0 5/5 a.v.
locality 41

CASG 66255.01 ale 16.0 132 15.6 3/5 a.v.

CASG 66255.03 LES 16.5 16.6 14.0, 3/5 a.v.

CASG 66255.04 14.1 16.5, 14.8 14.3 3/5 a.v.

CASG 66255.05 14.2 18.1 14.5 16.8, 3/5 ay.

CASG 66255.06 19.7 21.6 15.0 18. 3/5 a.v.

CASG 66255.02 18.3. 18.8. 19.7, 24.0. 3/5 a.v.
locality 42

CASG 66256.01 Ud 10.5, Sie 8.4 3/4 a.v.
locality 43

CASG 66257.01 13.8 16.9 14.0 14.0 3/5 a.v.

CASG 66257.02 14.8 17.6 17.0 16.6 3/5 a.v.

Formation, Triassic cover on the Golconda Allo-
chthon, zones LNo2-LNo3 of Tozer (1980), based on
stratigraphic position], 42 [Nevada, Dun Glen For-
mation, Triassic cover on the Golconda Allochthon,
zone LNo3 of Tozer (1980), based on associated fau-
na], and 43 [Nevada, ?Winnemucca Formation, Tri-
assic cover on the Golconda Allochthon, zones LNo2-
LNo3 of Tozer (1980), based on stratigraphic position].
It also appears to be present in rocks assigned to the
““Middle Triassic’ of Peru by Tilmann (1917), where
it was reported as Spiriferina sp. The dorsal valve ex-
terior figured by Tilmann (1917, pl. 25, fig. 12a), shows
the trifurcate costae on the fold that are characteristic
of this species.

Diagnosis.—Paucicostate Spondylospira bearing an
odd number of costae on the dorsal fold, which arise
from bilateral trifurcation of a single initial mesial cos-
ta.

Types.—Holotype, CASG 66247.01; paratypes,
CASG 66247.02—CASG 66247.04; figured specimens,
CASG 66255.01, CASG 66255.02, CASG 66256.01,
USNM 450289; measured specimens, CASG
66255.01-CASG 66255.06, CASG 66256.01, CASG
66257.01, CASG 66257.02, USNM 450288.

Comparison.—S. tricosta, n. sp. may be distin-
guished from most species of Spondylospira by its pau-
cicostate macro-ornament. Among paucicostate cyr-
tinoid spiriferinacean species, it may be distinguished
from SS. /ewesensis (Lees) by the odd number of costae
borne on its dorsal fold. It may be distinguished from

Vitimetula parva, n. sp. by its more pronounced or-
nament, and presence of a dorsal fold. It may be dis-
tinguished from species of Zugmayerella Dagys, 1963,
which it resembles in form and outline, by its costate
fold and sulcus.

Discussion.—S. tricosta, n. sp. is probably closest in
overall appearance to S. /ewesensis, although no tran-
sitional forms have been observed. As is the case in
S. lewesensis, sporadic bifurcation of mesial and lateral
costae results in both paucicostate and multicostate
phenotypes, although the transition from one to the
other is not so well-represented by samples as is the
case with S. /ewesensis. Samples of S. tricosta from
Nevada most resemble species of Zugmayerella Dagys,
1963, differing most obviously in bearing costae on the
fold and sulcus.

Genus VITIMETULA, new genus

new genus aff. Spondylospira. Hoover, 1983, p. 1026.

Etymology of name.—L. vietus = wrinkled + L. me-
tula = conical or pyramidal figure.

Description.—Strongly inequibiconvex shells with
high spire-like ventral and shallow bowl-like dorsal
valves. Both valves may bear faint simple costae; cos-
tae more pronounced distally. Distinct, shallow, very
narrow ventral sulcus, extending from apex to anterior
commissure; no corresponding dorsal fold.

Ventral valve outline variable in ventral aspect. In-
terarea margins laterally rimmed; interarea surface pla-
nar, bearing coarse, sinuous vertical ridges, which may

86 BULLETIN 337

bifurcate commissurally, ridges terminating at hinge
in prominent simple denticles. Delthyrium high, nar-
row, open. Apical pedicle foramena multiple, “hon-
eycombed” as typical of subfamily.

Dorsal valve quite shallow, commonly widest at
hinge. Surface smooth or distally faintly costate.

Ventral interior with shallow spondylium. Articu-
lation by hinge denticle and by anterior portions of
dental plates that apparently match hinge sockets of
opposite valve. Surface smooth or faintly reflecting
exterior ornament. Muscle scars indistinct.

Dorsal interior posteriorly bearing short articulatory
shelf; shelf bearing anteroposteriorly elongate fossae
that accomodate articulatory hinge denticles of op-
posite valve. Shallow hinge sockets present. Median
septum absent, but low median myophragm separating
muscle scars. Umbo poorly developed, not produced
posterior to hinge line. Descending branches of spire
produced anteriorly from anterior margins of hinge
sockets, anteriorly forming a jugum. All elements joined
to valve floor by complex meshwork.

Type species.— Vitimetula parva, n. sp.

Diagnosis.—Faintly costate Spondylospirinae with
high ventral and shallow dorsal valves, narrow shallow
ventral sulcus but no dorsal fold. Interior as for Spon-
dylospira Cooper.

Occurrence. — Vitimetula, n. gen. is at present known
only from locality 15 (Idaho, Wallowa terrane), where
it co-occurs with Spondylospira lewesensis (Lees, 1934).
The fauna at this locality is probably of late Norian
age (Stanley, 1979) [zones UNol-UNo3 of Tozer
(1980). See Appendix for discussion of problems with
the age of this locality.].

Comparison.—Vitimetula, n. gen. is easily distin-
guished from Spondylospira by the absence of a dorsal
fold and the absence of costae on the mesial portions
of either valve. It is distinguished from other laballids
and spondylospirids that have smooth mesial portions
by its narrow sulcus and comparatively smooth surface
overall. It is distinguished from Psioidea Hector, 1879
by the absence of a dorsal fold and its relatively short
hingeline and non-transverse outline.

Discussion.—A single specimen of the type lot of
Spondylospira reesidei Cooper [USNM 103468d] is here

issigned to Vitimetula, n. gen.

Vitimetula parva, new species
Plate 11, figures 16-32

reesidei Cooper, 1942, p. 232 [part: unfigured para-
fi 1M 103468d)]; Cooper, in Shimer and Shrock, 1944, p.
> [part: unfigured paratype (USNM 1034684d)].

Description.—Small, strongly inequibiconvex shells
with high conical ventral and shallow bowl-like dorsal
valves. Both valves bearing faint simple costae; costae

more pronounced distally. Distinct, very narrow, shal-
low ventral sulcus, but no corresponding dorsal fold.

Ventral valve commonly four times as deep as dor-
sal; outline in ventral aspect variable. Margins of in-
terarea commonly produced posteriorly as low mim;
posterodorsal hinge margins commonly flared distally.
Interarea bearing coarse, sinuous, dorsally bifurcate
vertical ridges; ridges terminating at hinge in promi-
nent simple denticles. Apical portion of area pierced
by several posteroventrally oriented slitlike openings
(see Pl. 11, fig. 18). Delthyrium long, narrow, open.

Dorsal valve quite shallow, commonly widest at
hinge; surface smooth or marginally faintly costate.

Ventral interior with shallow spondylium formed by
junction of short dental plates and median septum;
septum length variable in terms of spondylium sup-
port, invariably projecting posteriorly into spondy-
lium. No functional paired hinge teeth, but anterior
portions of dental plates apparently articulating with
hinge sockets of opposite valve. Articulation mostly
by hingeline denticles. Muscle scars not evident. In-
terior surface smooth or faintly reflecting subdued ex-
terior costation.

Dorsal interior posteriorly bearing short articulatory
shelf. Shelf bearing anteroposteriorly elongate fossae
which accomodate articulatory hinge denticles of op-
posite valve. Shallow hinge sockets present. No median
septum, but low rounded myophragm separating stri-
ate muscle scars. Umbo poorly developed, not pro-
duced posterior to hinge line. Descending branches of
spire produced anteriorly from anterior margins of
hinge sockets, anteriorly forming a jugum, all elements
joined to valve floor by complex meshwork.

For measurements, see Table 7.

Occurrence.— Vitimetula parva, n. sp. is at present
known only at locality 15 (Idaho: Wallowa terrane),
where it co-occurs with Spondylospira lewesensis (Lees,
1934). The fauna there suggests a late Norian age [Stan-
ley (1979); zones UNol-UNo3 of Tozer (1980), based
on associated fauna, but see Appendix for discussion
of problems with this age].

Diagnosis.—Small Vitimetula.

Types.—Holotype, USNM 450290; paratypes,
USNM 103468d, USNM 450291-USNM 450298.

Comparison.— Vitimetula, n. gen. is monotypic. V.
parva, n. sp. may be distinguished from species of
Spondylospira by its smooth narrow ventral sulcus and
its lack ofa dorsal fold, and from other spondylospirids
and laballids by its high ventral valve, smooth exterior,
and narrow ventral sulcus. Its internal details indicate
that it is probably most closely related to Spondylos-
pira. Its ventral valve somewhat resembles that of Zug-
mayerella americana, n. sp., but consistently bears a
narrower sulcus, while the dorsal valves of the two
forms are markedly dissimilar.

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 87

Table 7.—Measurements (in mm) of type specimens of Vitimetula parva, n. gen. and n. sp. See p. 74 for explanations of abbreviations and

subscripts.
height/depth or
hinge width maximum width length thickness specimen type

locality 15

. USNM 450290 (holotype) 4.8 6.3 6.0 10.2 a.v.
USNM 450291 (paratype) 5:5 5:5 8.2 6.2 v.V.
USNM 450292 (paratype) 5.4 5.4 3.8 4.5 V.V.
USNM 450293 (paratype) 6.04. 6.05. SiS 6.3 v.V.
USNM 450294 (paratype) BiOne 3/6 3.7 4.2 V.v.
USNM 450295 (paratype) 5:5 5:5 5.0 1.0 d.v.
USNM 450296 (paratype) 7.0 7.0 4.8 135. d.v.
USNM 450297 (paratype) 6.0, Ue. 4.5. Dis d.v.
USNM 450298 (paratype) 9.0, 9.0, => 1.6 d.v.
USNM 103468d (paratype) 8.0 8.0 6.0. 4.8 V.V.

Family LABALLIDAE Dagys, 1962
Subfamily LABALLINAE Dagys, 1962
Genus ZUGMAYERELLA Dagys, 1963

Type species.—Spiriferina koessenensis Zugmayer,
1882, p. 28, pl. 3, figs. 2, 3, 5, 13.

Diagnosis.—Small to medium-sized laballines with
high ventral valve, smooth, sharply differentiated fold
and sulcus, and paucicostate flanks. Interior as for
Spondylospira Cooper, 1942. Microornament of
densely packed spines.

Occurrence.— Zugmayerella has been recovered from
Norian strata in the Alps, the Carpathians, the Balkans,
the Dinarides, Crimea, the Caucasus, northeastern
U.S.S.R., Papua New Guinea, China, Tibet, New
Zealand, and South America. In the present collec-
tions, Zugmayerella and a species questionably re-
ferred to that genus have been recovered from seven
localities in Alaska, California, and Oregon, ranging in
age from late Karnian to early Norian.

Comparison.— Zugmayerella may be distinguished
externally from Spondylospira Cooper, 1942, by its
smooth, rather than costate fold and sulcus. It differs
from Vitimetula, n. gen. in its more transverse outline,
broader sulcus, and presence of a dorsal fold. It differs
from Spinolepismatina Dagys, 1974, and Pseudola-
balla Dagys, 1974, in having a longitudinally ridged,
rather than smooth interarea. It differs from K/ipstei-
nella Dagys, 1974, in its denticulate hinge, and absence
of a deltidium, and from Laballa Moisseiev in Dagys,
1962, by its denticulate hinge and by the strong pli-
cation of both valves. It differs from Orientospira Da-
gys, 1965, in its subpyramidal rather than globose shell
form, and its comparatively high, narrow interarea. It
differs from Phenacozugmayerella, n. gen. in its spi-
nose, rather than capillate—cancellate microornament.

Discussion.— Placement of the genus Zugmayerella
in the Laballidae rather than the Spondylospiridae is
tentative. It was so assigned by Dagys (1962), and the
critical characters of the ventral apex necessary to dis-

tinguish between the two families are not well-shown
in any of the specimens of Zugmayerella available for
this study. It is, however, possible that better preser-
vation in specimens recovered in the future may dictate
that this genus be placed in the Spondylospiridae with
the other genera described in this paper. Zugmayerella
was originally distinguished from Lepismatina Wang,
1955 (Dagys, 1963, pp. 99-100) on the basis of its
ridged interarea and resultant denticulate hinge, fea-
tures which Dagys believed were not shared by Lep-
ismatina. Later examination by Sun (1981) of topo-
typic material of MLepismatina revealed that
Lepismatina does not belong in the Laballidae, but
rather in the Spiriferinidae.

One character common to known species of Zug-
mayerella is spinose microornament (Dagys, written
commun., 1990). This character, though mentioned in
the original diagnosis of the genus (Dagys, 1963), was
not there accorded the importance brought out by later
observations (Dagys, written commun., 1990). The
spines in Zugmayerella are small and easily broken,
and a pustulose surface microornament is commonly
the result. Pearson (1977, p. 23), in one of the few
comprehensive discussions of the genus in English,
states that the shell surface is “‘pustulose, possibly spi-
nose.” It is this character that, among others, sets this
genus apart from the new genus Phenacozugmayerella,
in which the cancellate microornament is formed by
beaded capillae.

Zugmayerella americana, new species
Plate 11, figures 33-50; Plate 12, figures 1—5

Description.—Small, unequally biconvex shells hav-
ing high, pyramidal ventral and shallow bowl-like dor-
sal valves. Ventral sulcus and dorsal fold only slightly
developed. Microornament of densely-packed spines.

Ventral valve semicircular in ventral aspect, trian-
gular in anterior aspect. Depth commonly subequal to
shell width, slightly less than height. Median sulcus
shallow, rounded, bounded laterally by large plicae.

88 BULLETIN 337

Table 8.—Measurements (in mm) of type specimens of Zumayerella americana, Nn. sp. See p. 74 for explanations of abbreviations and

subscripts.
ee
hinge width maximum width length thickness specimen type
locality 3
USNM 450305 (paratype) 5.8), 5.8), 4.1 4.0 V.V.
locality 20
USNM 450299 (paratype) 4.8, 6.0, 4.1 3.9 V.V.
USNM 450300 (paratype) 6.2 6.8 4.4 4.54. V.V.
USNM 450301 (paratype) 11.8 13.2 10.0 9.1 V.V.
USNM 450302 (paratype) 10.5 14.1 10.3 3.4 d.v.
USNM 450303 (paratype) 8.8 13.3 1253 3.2 dv.
locality 21
USNM 450304 (paratype) Wes) 8.4 6.5 4.3 V.V.
locality 28
USNM 450306 (holotype) 5.4 5.4 51 4.4 a.v.
USNM 450307 (paratype) 7.9, TES 4.6 4.7 V.V.
USNM 450308 (paratype) 4.6 7.6 6.1 4.6 V.V.
USNM 450309 (paratype) 9.6, 12.0 10.0 9.8 V.V.
USNM 450310 (paratype 122) 14.1 10.0 9.7 V.v.
USNM 450311 (paratype) 10.0 11.3 7.6 3.2 dv.
USNM 450312 (paratype) 7.9. 11.6 8.8 2.3 d.v.
USNM 450313 (paratype) 10.6 11.6 10.0 2.6 d.v.

Flanks bearing four to six prominent plications. In-
terarea sharply recurved posteriorly at apex, otherwise
flat; entire surface vertically ridged, ridges terminating
as denticles at hinge. Delthyrium open, narrow. Hinge
denticulate; primary teeth absent, anterior portions of
dental plates may be marginally functional in articu-
lation.

Dorsal valve quadrate to transverse or semicircular
in dorsal aspect, broadly rounded in anterior or lateral
aspects. Umbo tiny, not produced posterior to hinge-
line. Median fold low, square-shouldered, medially de-
pressed; bordered laterally by four to six prominent
rounded plications; fold commonly lower than adja-
cent plicae. Plicae originating from central portion of
hingeline, but not from mesially-located “point-source”
beak.

Ventral interior generally reflecting exterior macro-
ornament. Thin median septum extending one-half to
two-thirds valve length, may be asymmetric with re-
spect to sulcus. Dental plates united with median sep-
tum just below its crest, forming small spondylium.
Muscle scars indistinct.

Dorsal interior with simple, anteroposteriorly striate

| process. Fossae accomodating denticles of

ngeline anteroposteriorly elongate, may be

1esially. Hinge sockets present, probably

nally functional in articulation. Low median my-

phragm present or absent, if present extending about

me-half valve length. Descending branches of spire

joined to valve floor at about mid-length of valve,
united anteriorly to that point by simple jugum.

For measurements, see Table 8.
Diagnosis.—Small Zugmayerella with pyramidal,

comparatively high ventral valve, shallow, flat-bot-
tomed ventral sulcus and low, mesially depressed dor-
sal fold, and four to six lateral plications. Dorsal umbo
not produced posterior to hingeline.

Occurrence.— Z. americana, n. sp. has been recov-
ered in this study from rocks ranging in age from mid-
dle late Karnian [zone UC2 of Tozer (1980)] to earliest
Norian [zone LNol of Tozer (1980)], at localities 3
(California, 7Hosselkus Formation, Eastern Klamath
terrane, zone UC2 of Tozer (1980), based on asso-
ciated fauna], 13 (Alaska, Wrangellia terrane, age un-
known), 20 (Alaska, beds transitional between Chiti-
stone and Nizina formations, Wrangellia terrane, zone
LNol of Tozer (1980), based on associated fauna], 21
[Oregon, Martin Bridge Formation, Wallowa terrane,
zone UC3 of Tozer (1980), based on associated fauna],
and 28 (Oregon, Martin Bridge Formation, Wallowa
terrane, zone LNol of Tozer (1980), based on asso-
ciated fauna]. All specimens recovered are silicifed.
The fine preservation of this form wherever it is found
and its comparatively wide distribution, coupled with
its relatively short stratigraphic range, combine to make
it a fine potential biostratigraphic index. Etching of
material in suitable matrix should provide usable spec-
imens of this small but important form.

Types.—Holotype, USNM 450306; figured para-
types, USNM 450300-450302, USNM 450305,
USNM 450308, USNM 450309, USNM 450311,
USNM 450313; measured paratypes, USNM 450299-
USNM 450313.

Comparison.— Zugmayerella americana, n. sp. may
be distinguished from most known species of the genus
by its lower fold and shallower sulcus, and by lacking

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 89

a dorsal umbonal bulge that projects posterior to the
hingeline. It is smaller and less globose than Z. eurea
Dagys, 1965, Z. inaequiplicata Dagys, 1965, or Z.
koessenensis Zugmayer, 1882. It is smaller and has
more lateral plications than does Z. hemipyramida Sun,
1981, and is smaller, and has plications that are less
angular than those of Z. uncinata (Schafhautl, 1851).
It is smaller, narrower, and lacks the characteristic rib
in the sulcus and furrow on the fold that are charac-
teristic of Z. osmana (Bittner, 1902).

Among the other North American cyrtinoid spiri-
ferinaceans, it is easily distinguished from specimens
of Phenacozugmayerella mimuncinata, n. sp., by its
smaller size and less acute plications and by the lack
of spinose ornament in the latter form. From all the
species of Spondylospira Cooper, 1942, and from Pseu-
dospondylospira perplexa, n. gen. and n. sp., Dagys-
pirifer fascicostata, n. gen. and n. sp., and Vitimetula
parva, n. gen. and n. sp., it is distinguished by its ap-
parent lack of pedicle foramena in the ventral valve.

The small shell of Zugmayerella americana, n. sp.
is most similar to Z. yueliangpingica Ching and Feng,
1977 from the Upper Triassic of Gueizhou, China,
from which it differs in size. The latter form, described
on the basis of only four specimens, two ventral ex-
ternal molds, and one dorsal external mold, and one
dorsal internal mold, ranges from 15 to 20 mm wide,
while the largest known specimen of Z. americana is
just over 14 mm wide.

Discussion.— Zugmayerella americana, n. sp. be-
longs to the only cosmopolitan cyrtinoid genus among
those described in this report. Its close similarity to
the Chinese Z. yueliangpingica Ching and Feng, 1977
may suggest origin via trans-Pacific dispersal.

Zugmayerella species
Plate 12, figure 34

Discussion.—A single silicified dorsal valve [USNM
450317] assignable to Zugmayerella, but not confi-
dently associated with any species, shows the sup-
porting structures of the brachidium in best detail. The
descending branches of the spire extend anterodorsally
from the hingeline and contact the valve floor at about
midlength. A simple jugum unites the branches ante-
riorly. It was recovered from locality 1 [California,
?Hosselkus Formation, Eastern Klamath terrane, zones
UCI-LNo2 of Tozer (1980), based on stratigraphic
position].

Measurements (in mm)*.— Hinge width, 10.6,,.; valve
width, 10.6,. ; valve length, 8.8,..; valve height, 5.0.

Type.— Figured specimen, USNM 450317.

*see p. 74 for explanations of abbreviations and subscripts.

Table 9.— Measurements (in mm) of specimens of Zugmayerella?
sp. A. See p. 74 for explanations of abbreviations and subscripts.

maxi- speci-
hinge mum men
width width length height type
locality 3

USNM 450315 122 17.6, 15. — V.V.

locality 12
USNM 450314 10.5 14.2 12.7 11.6 V.V.
USNM 450316 7529 24315 13.7, IWS 2 V.V.

Zugmayerella? species A
Plate 12, figures 25-33

Discussion.— Three partial ventral valves (one silic-
ified, two calcareous) of a new species here question-
ably assigned to Zugmayerella were recovered from
localities 3 [California, 7Hosselkus Formation, Eastern
Klamath terrane, zone UC2 of Tozer (1980), based
on associated fauna] and 12 (Oregon, ?Martin Bridge
Formation, Wallowa terrane, age unknown). All ap-
pear to have been somewhat decorticated, but repre-
sent a pauciplicate form with a smooth broad sulcus
and very faint lateral plications. The rarity and the
poor preservation of this form make formal species
designation unwarranted here, and the lack of un-
equivocally preserved microornament make generic
identification questionable.

For measurements, see Table 9.

Types.— Figured specimens, USNM 450314, USNM
450315; measured specimens, USNM 450314-USNM
450316.

Genus PHENACOZUGMAYERELLA, new genus

Etymology of name.—Gr. phenax = cheat, impostor,
+ Zugmayerella.

Type species.— Phenacozugmayerella mimuncinata,
Nn. sp.

Diagnosis.— Medium-sized laballines with high ven-
tral valve, smooth, sharply differentiated fold and sul-
cus, narrow denticulate hinge, and paucicostate flanks.
Surface microornament cancellate, with capillae bro-
ken into transverse bars by growth increments (Text-
fig. 5).

Occurrence.—As for species.

Comparison. — Phenacozugmayerella is perhaps most
easily confused with Zugmayerella Dagys, 1963, from
which it differs in its microornament, which is capil-
late-cancellate rather than densely spinose. In this re-
spect is also differs from most other laballids, except
aspinose forms like Yanospira Dagys, 1977, which dif-
fers in having a duplex interarea, with the central por-
tion ridged, and the outer part smooth, and by being
more transverse. It may be distinguished from Spon-
dylospira Cooper, 1942, by its smooth, rather than

90 BULLETIN 337

Table 10.—Measurements (in mm) of type specimens of Phenacozugmayerella mimuncinata, n. gen. and n. sp. See p. 74 for explanations

of abbreviations and subscripts.

eS

maximum — dorsal valve

hinge width width

locality 31
CASG 66248.03 (paratype) 14.5 19
CASG 66248.04 (paratype) 19 21.5
CASG 66248.06 (paratype) 13 15
locality 32
CASG 66249.01 (paratype) 9.5 9.5
CASG 66249.02 (paratype) 11.5 13.5
locality 34
CASG 66250.01 (paratype) 15 17
CASG 66250.02 (holotype) 18.5 20
locality 35
CASG 66251.02 (paratype) 19 19
CASG 66251.03 (paratype) 17 20
locality 36
CASG 66252.01 (paratype) 20 21

interarea specimen
length total length __ thickness height type
15 19.5 18.5 11.5 a.v.
15 20 18 12 a.v.
10 12.5 13. 10.5. a.v.
5.5 Te 6.5 6.0. a.v.
10.5 10.5 11.0, 10.0. a.v.
16 21 17 9 a.v.
12.5 DAES 17 12.5 a.v.
14 17 15.5 10 a.v
18.5 20. 19 12. d.v
15 18.5 16.5 13 a.v

Se ee eee eee

costate fold and sulcus, and from Vitimetula, n. gen.,
Dagyspirifer, n. gen., and Pseudospondylospira, n. gen.,
by its equidimensionally pyramidal plicate shell, and
from all of these by its apparent lack of a functional
pedicle.

Discussion.— Assuming (with Dagys, 1974) that the
nature of the microornament in cyrtinoid spiriferina-
ceans is relatively important in distinguishing among
the various genera, Phenacozugmayerella is probably
most closely related to Yanospira Dagys, 1977, from
the Norian of Okhotsk, U.S. S. R.

Phenacozugmayerella mimuncinata, new species
Plate 12, figures 6-24

Zugmayerella uncinata (Schafhautl). Stanley, 1979, p. 14 [as Z. in-
cinata], pl. 8, figs. 10-12] (non Spirifer uncinata Schafhautl, 1851,
p. 135, pl. 24, fig. 33).

Etymology of name.—Gr. mimos = actor + uncin-
ata.
Description.—Shell small- to medium-sized, un-
equally biconvex, with pronounced dorsal fold and
ventral sulcus. Fold height about two to four, com-
monly two, times the height of adjacent plicae. Flanks
f both valves corrugated by radial plicae that expand
plicae of less magnitude laterally. Fold and
to subrounded in section; commonly
iced, rarely interrupted mesially by one or
) faint radial plicae. Valve outline in ventral aspect
juadrate, with straight lateral margins that converge
slightly anteriorly, and mesially incurved anterior mar-
zin. Outline in dorsal aspect transverse with rounded
anterior margin in small specimens, becoming quad-
rate in larger, apparently more mature individuals.
Outer shell layer commonly not preserved; where pre-

served, surface micro-ornament of radial capillae,
composed of obliquely-disposed transverse bars formed
by the intersection of capillae with growth lines, the
two elements of microornament together presenting a
finely cancellate surface. Three to five, most commonly
three plicae on either side of fold on dorsal valve, one
or more on ventral. Hinge width subequal to maximum
shell width.

Ventral valve deep, varying from about one-half to
four-fifths of hinge width in height. Valve length and
width, and shell thickness all subequal, but variable.
Interarea high, triangular, with narrow to moderately
broad triangular delthryium; interarea surface longi-
tudinally ridged, ridges fine, extending dorsal to hinge
as fine denticles; denticles fitting into articulatory re-
cesses along hinge of opposite valve. Interarea flat,
catacline to apsacline, angle to commissural plane mea-
sured through shell 90° to about 110°, most commonly
about 100°. Ventral apex of ventral beak commonly
hooked posteriorly in last few mm.

Dorsal valve comparatively shallow, with fold pro-
viding curved quadrate section in lateral view. No in-
terarea exposed, but short low beak extending about |
mm beyond the hinge line in well-preserved speci-
mens. Lateral plicae not straight, but gently curved and
concave distally.

Ventral interior with strong median septum extend-
ing along valve floor about one-half valve length, rising
anteriorly as a thin blade. Apparent dental plates join-
ing inner margins of delthyrium to about mid-height
of median septum, to form spondylium; median sep-
tum continuing posterior to spondylium as thin blade
not quite reaching plane of interarea surface. Muscle
scars indistinct. Interior surface plicate, reflecting ex-
terior macro-ornament.

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 91

Dorsal interior with minute notothyrium and hinge-
line bearing row of anteroposteriorly elongate fossae
that accommodate ventral valve hinge denticles.

For measurements, see Table 10.

Occurrence.—Phenacozugmayerella mimuncinata,
Nn. sp. 1s apparently endemic to North America. In the
present collections it has been recovered from rocks
ranging in age from late Karnian to middle Norian
[zones UCI through MNo2 of Tozer (1980)], at local-
ities 6 [Alaska, Alexander terrane, zones UC1—UC3 of
Tozer (1980), based on associated fauna], 32 [Nevada,

Osobb Formation, Triassic cover on the Golconda Al-
lochthon, zones LNol-LNo3 of Tozer (1980), based
on stratigraphic position], 33 [Nevada, lower member
of Luning Formation, Paradise terrane, zone LNol of
Tozer (1980), based on fauna at this stratigraphic level],
34 [Nevada, upper member of Luning Formation, Par-
adise terrane, zone LNo3 of Tozer (1980), based on
stratigraphic position], 35 [Nevada, Luning Forma-
tion, Paradise terrane, zone LNo3 of Tozer (1980),
based on associated fauna], 36 [Nevada, Luning For-
mation, Paradise terrane, zone LNol1 of Tozer (1980),

Text-figure 5.—Posterodorsolateral views of portions of a dorsal valve of Phenacozugmayerella mimuncinata, n. gen. and n. sp., CASG
66250.02 (holotype, from loc. 36), showing (a) the five folds on one flank, <8; (b) the surface microornament of beaded capillae on the next-
to-the-top fold, x 17; and (c) the capillate and capillate—cancellate (x) surface microornament on the top fold.

92 BULLETIN 337

based on fauna at this stratigraphic level], 37 [Nevada,
lower Luning Formation, Paradise terrane, zones
LNol-LNo2 of Tozer (1980), based on fauna at this
stratigraphic level], 38 [Nevada, Luning Formation,
Paradise terrane, zones LNol-MNo2 of Tozer (1980),
based on stratigraphic position], 45 [Nevada, Jungo
terrane, zone MNo2 of Tozer (1980), based on asso-
ciated fauna], 48 [Oregon, Brisbois Formation, Izee
terrane, zones UC1—UC3 of Tozer (1980), based on
stratigraphic position], 49 [Oregon, Brisbois Forma-
tion, Izee terrane, zones UC1I—UC3 of Tozer (1980),
based on stratigraphic position], and 51 [Oregon, Bris-
bois Formation, Izee terrane, zones UC1—UC3 of Toz-
er (1980), based on stratigraphic position].
Diagnosis.—Small to medium-sized pauciplicate

Phenacozugmayerella with acute-angled plications and
distinct, comparatively large fold and sulcus.

Types.—Holotype, CASG 66250.02; figured para-
types, CASG 66248.03—CASG 66248.05, CASG
66249.01, CASG 66249.02, CASG 66250.01, CASG
66251.02, CASG 66252.01; measured paratypes,
CASG 66248.03, CASG 66248.04, CASG 66248.06,
CASG 66249.01, CASG 66249.02, CASG 66250.01,
CASG 66251.02, CASG 66251.03, CASG 66252.01.

Comparison. — Phenacozugmayerella mimuncinata,
n. sp. is the only species of the genus, and is most easily
confused with Zugmayerella uncinata (Schafhautl,
1851), from which it differs in its capillate-cancellate
surface micrornament, and in lacking the duplex in-
terarea that is characteristic of that species.

APPENDIX

LOCALITIES CITED IN THIS REPORT

Locality information from specimen labels (not enclosed in square brackets) in many cases has been enhanced
by comparison of original register entries, field labels, maps, notebooks, more recent topographic maps, and
geologic maps and reports. Such additions appear in brackets. Zonal abbreviations refer to the zonation for the
Triassic presented by Tozer (1980, Table II; see Table 1 herein) and have been assigned by N. J. Silberling (written
commun., 1989). Terrane assignments were made by comparison of map position with those in the “Folio of the
Lithotectonic Terrane Maps of the North American Cordillera” [U. S. Geological Survey, Miscellaneous Field
Studies Maps, MF-1874-A (Jones et al., 1987), MF-1874-B (Monger and Berg, 1987), and MF-1874-C (Silberling
et al., 1987)], and by consultation with N. J. Silberling (written commun., 1989). Locality numbers as used in
this report (see also Text-fig. 1) are followed by the field number, in parentheses.

Locality 1 (USGS loc. 2309).— California, [Shasta Co., Bollibokka
Mtn. 15’ quad.;] E. line of sec. 30, T. 34 N., R. 1 W.; on both sides
of Little Cow Creek [= Little Cedar Ck.], just below mouth of Bear
Gulch; Collectors: [J.] Storrs and [C.] Washburne, August 8, 1901.
[formation not given, probably Hosselkus Fm.; zones UCI-—LNo2
of Tozer (1980), based on stratigraphic position; Eastern Klamath
terrane].

Locality 2 (USGS loc. 2314).—California, Shasta Co., [Burney 30’
quad.] near ctr., sec. 20, T. 34 N., R. 1 W.; 5 mi NE of Furnaceville;
Collectors: [J.] Storrs and [C.] Washburne, August 8, 1901. [for-
mation not given, probably Modin Fm.; zones MNol-UNo2 of
Tozer (1980), based on associated fauna; Eastern Klamath terrane]

Locality 3 (USGS loc. 2446).—California, Shasta Co., [?Millville
15’ quad.] on Toll Road by Cedar Creek, 10'2 mi [?] below [west of]
Round Mtn.; Collectors: J. S. Diller and J. Storrs, June, 1892. [for-
mation not given, probably Hosselkus Fm.; about zone UC2 of Tozer
(1980), based on associated fauna; Eastern Klamath terrane]

Locality 5 (USGS loc. 2769).— California, Shasta Co., [?Big Bend
15’ quad.] 5 mi. E. of Grizzly Peak, W. face of Table; bed of s. s.

d cong.; Collector: J. Storrs, June 19, 1903. [Formation not given,

lin Formation (based on lithology); zones LNol1-UNo3 of Toz-
based on associated fauna; Eastern Klamath terrane]
SGS loc. 10548).— Alaska, Southeast, Screen Islands,
3} 1:63,360 quad.;] Clarence Strait, west coast of Etolin
35’ of dark gray limestone 400’ thick; Collector: E. M.
e, 1905. [Formation not given; zones UC1—UC3 of Tozer (1980),
n associated fauna; Alexander terrane]

’ (USGS loc. 11406).—Alaska, Southeast, [Port Alex-
ander D—1 1:63,360 quad.]; west side of Keku Straits, 7 mi NW of
Pup Island, Kuiu Islands; Collector: A. F. Buddington, 1922. [Corn-
wallis Limestone; zones UC1—LNol(?) of Tozer (1980), based on
associated fauna (Muffler, 1967, p. C34); Alexander terrane]

Locality 8 (USGS loc. 11814).—Nevada, [Mineral Co.,] Haw-
thorne [1°] quad., T. 7 N., R. 35 E., N. 65° E. from Mina, first lime-
stone exposure in Dunlap Gulch; Collectors: [H. G.] Ferguson, [S. H.]
Cathcart, Heiges and [T. W.] Stanton, July 13, 1923. [Lower member
of the Luning Formation, zone LNol of Tozer (1980), based on
associated fauna (sponges); Paradise terrane]

Locality 10 (USGS loc. 13354).—Nevada, [Mineral Co., Haw-
thorne 1° quad.;] Gabb’s Valley Range, S. of Mayflower Ball Mill E.
of Road, 100'—200' above top of blue limestone with small amount
of brown sandy material; Collector: H. G. Ferguson, September 2,
1925. [Middle member of Gabbs Formation, zones UNo2-UNo3
of Tozer (1980), based on associated fauna; Paradise terrane]

Locality 12 (USGS loc. 15550).—Oregon, [Wallowa Co.,] Imnaha
River, [Harl Butte 15’ quad.;] NE, sec. 15, T. 2 S., R. 48 E., isolated
outcrop over more or less 2 square mile area; Collectors: [R. W.]
Richards and [B. N.] Moore, August 14, 1930. [formation not given,
??Martin Bridge Fm.; no information on stratigraphic position; Wal-
lowa terrane]

Locality 13 (USGS loc. 16266).—Alaska, South Central, Nutzotin
Mtns., [Nabesna C-5 1:63,360 quad.;] Long. 143°10' W.; Lat. 62°35’
N.; head of tributary of Jack Creek that lies directly S. of Bear Ck.
glacier, Copper River Region; Collector: F. H. Moffit, August 30,
1931. [formation not given; Wrangellia terrane]

Locality 15 (USGS loc. 17424).—Idaho, [Nez Perce-Lewis Co.
line, Culdesac 15’ quad.;] about 6 mi up Mission Ck. from Jaques
R. R. Station, past St. Joseph’s Mission and Slickpoo Settlement, at
first narrows of Mission Creek, collection from black limestone abt.
70' above water, E. side of ck.; Collectors: J. Reed and J. S. Williams,
June 17, 1936. [formation not assigned, zones UNol-UN0o3 of Tozer
(1980), based on associated fauna [the brachiopods (Cooper, 1942)
and gastropods (Haas, 1953) suggest a Norian—Rhaetian age. Stanley
(1979) notes that many of the coral species are not known elsewhere

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 93

in North America, but have been reported from the Zlambach beds
(Norian) of Fischerweise, Austria. He states (p. 27) that he has “‘cho-
sen tentatively to regard the Lewiston locality as Upper Norian,”
but that “no diagnostic ammonites have been discovered, and the
age of the known fauna is equivocal.”; Wallowa terrane]

Locality 16 (USGS loc. 24387).—Peru, Atacocha Dist.; Dept. of
Pasco; Machican Mine area; Collectors: R. F. Johnson and R. W.
Lewis, 1952. [formation not given, = Chambara Fm. of Pucara Gp.;
zones UC1-—UNo3 of Tozer (1980), based on associated fauna; ter-
rane unknown]

Locality 18 (USGS loc. M96).—Nevada, Pershing Co., [Buffalo
Mtn. 15’ quad.;] NE", sec. 5, T. 26 N., R. 34 E, and sec. 32, T. 27 N.,
R. 34 E.; along strike of beds at base of limestone cliffs, approx. 74
miS. to 2 mi W. of Nevada Quicksilver [= Juniper] mine; Collectors:
N. J. Silberling, R. E. Wallace, W. P. Irwin, 1956. [formation not
given, but = Antelope Springs Fm. of Oldow, Bartel, and Gelber
(1990); zone LNo3 of Tozer (1980), based on associated fauna; Jungo
terrane]

Locality 20 (USGS loc. M1708).—Alaska, Wrangell Mtns., Mc-
Carthy B—5 1:63,360 quad.; on west side of East Fork Valley just E.
of largest patch of Cretaceous capping Green Butte Ridge; Collector:
N. J. Silberling, July 7, 1962. Transitional beds between Chitistone
and Nizina formations. [zone LNol of Tozer (1980), based on as-
sociated fauna; Wrangellia terrane]

Locality 21 (USGS loc. M1749).—Oregon, [Wallowa Co.,] Wal-
lowa Mtns., Enterprise 15’ quad.; ctr. of W'2 of boundary line bet.
sec. 3, T. 3 S.,R. 44 E., and sec. 34, T. 2 S., R. 44 E.; elev. ca. 6240’
on W. side of Hurricane Ck., about 1 mi N. of Little Granite Ck.;
Collector: B. Nolf [Oregon State Univ.], 1962. Martin Bridge For-
mation. [zone UC3 of Tozer (1980), based on associated fauna;
Wallowa terrane]

Locality 22 (USGS loc. M1906).— Alaska, Southeast, Keku Strait,
Port Alexander [D-1 1:63,360] quad.; NE side of Cornwallis Pen-
insula ca. 1 mi E. of Point 5600’ S. 85° E. from triangulation station
Corn; silicified fauna in medium to thick-bedded limestone; Collec-
tor: L. J. P. Muffler, June, 1963. Cornwallis Limestone. [Silberling,
written commun., 1963 (cited by Muffler, 1967, p. C30, table 2)
dates this locality as of “‘uncertain age in the Late Triassic,” based
on “‘scleractinian corals and (or) the spiriferid brachiopod Spondy-
lospira; arcestid or clydoniacid ammonites.” He later (written com-
mun., 1989) assigned it to zones UC1—UNo of Tozer (1980), based
on associated fauna; Alexander terrane]

Locality 23 (USGS loc. M1911).— Alaska, Southeast, [Keku Strait,
Port Alexandria D-1 1:63,360 quad.]; S. side of small cove of E.
shore, Kuiu Island, 1.5 mi SSE, or 8300’ S. 19° E. from triangulation
station ‘“‘Low”; silicified fauna from 185’ above base of limestone;
Collector: N. J. Silberling, June, 1963. Cornwallis Limestone. [Sil-
berling, written commun., 1963 (cited by Muffler, 1967, p. C30,
table 2) dated this locality as “latest Karnian or earliest Norian,”
based on the occurrence of species of Halobia and Mojsisovicsites.
He later (written commun., 1989) assigned it to zone LNo1 of Tozer
(1980), based on associated fauna; Alexander terrane]

Locality 24 (USGS loc. M1917).—Alaska, Southeast, Keku Strait;
Port Alexander D-1 1:63,360 quad.; NE shore of Kuiu Island ca.
3% mi SE of Pt. Cornwallis; 17,950’ S. 69° E. from triangulation
station CORN; impure limestone 20-30’ above base of limestone;
Collectors: N. J. Silberling, L. J. P. Muffler, June 27, 1963. [Corn-
wallis Limestone; Silberling (written commun., 1963, cited by Muf-
fler, 1967, p. C28, table 1) dated this locality as of “‘uncertain age
within the Late Triassic,” based on “‘scleractinian corals and (or) the
spiriferid brachiopod Spondylospira, [and] arcestid or clydonitacid
ammonites’. He later (written commun., 1989) assigned it to zones
UCI-LNo2 of Tozer (1980), based on stratigraphic position; Al-
exander terrane]

Locality 25 (USGS loc. M1919).— Alaska, Southeast, Keku Strait,
Port Alexander D-1 1:63,360 quad.; E. shore Kuiu Island 0.5 mi

SSE, or 2500’ S. 23° E. from triangulation station LOW; impure
limestone interstratified with mafic tuff, but below main body of
Tmiassic volcanics; Collector: N. J. Silberling, June 29, 1963. Hound
Island Volcanics. [Silberling (written commun., 1963, cited by Muf-
fler, 1967, p. C30, table 2) dated this locality as “latest Karnian or
earliest Norian,”’ based on species of Halobia or Mojsisovicsites. He
later (written commun., 1989) assigned it to zones UC2-LNol of
Tozer (1980), based on associated fauna; Alexander terrane]

Locality 26 (USGS loc. M2135).— Alaska, Southeast, Keku Strait,
Port Alexander D-1 1:63,360 quad.; easternmost tip of islet in NW
Keku Islets, 4.41 mi S. 78.5° E. from triangulation station CORN;
dense thick-bedded limestone at tidal level; Collector: L. J. P. Muf-
fler, 1963. Keku Volcanics; Silberling (written commun., 1963, cited
by Muffler, 1967, p. C28, table 1) dated this locality as of “uncertain
age within the Late Triassic,” based on scleractinian corals and (or)
the spiriferid brachiopod Spondylospira, and arcestid or clydonitacid
ammonites. He later (written commun., 1989) assigned it to zones
LCI-—UN0o2 of Tozer (1980), based on associated fauna; Alexander
terrane]

Locality 27 (USGS loc. M2136).— Alaska, Southeast, Keku Strait,
Port Alexander D—-1 1:63,360 quad.; on S. side of islet in SE Keku
Straits, | mi SE of triangulation station THUM, 1.03 mi N. 34° E.
from triangulation station LOW; Collector: [L. J. P.] Muffler, 1963.
Cornwallis Limestone. [Silberling (written commun., 1963, cited by
Muffler, 1967, p. C30, table 2) dated this locality as of “uncertain
age in the Late Tnassic,” based on “‘scleractinian corals and (or) the
Spiriferid brachiopod Spondylospira; [and] arcestid or clydonitacid
ammonites.” He later (written commun., 1989) assigned it to zones
UCI1-UNo of Tozer (1980), based on stratigraphic position; Alex-
ander terrane]

Locality 28 (USGS loc. M2672).— Oregon, [Wallowa Co.,] Home-
stead 1:62,500 quad.; Oregon side of Snake River Canyon, elev.
2970’ on N. side McGraw Ck., abt. 0.85 mi NW from its mouth,
abt. 200’ about base of limestone unit; Collector: T. L. Vallier, June
25, 1964. [Martin Bridge Fm.; Silberling (cited in Vallier, 1967, pp.
246-247) assigned this locality to zone LNo1 of Tozer (1980), based
on the occurrence of the ammonoid Tropiceltites cf. T. columbianus;
Wallowa terrane]

Locality 29 (GSC loc. 10229).—Canada, Yukon Territory, Lake
Laberge area ca. 61° N. lat.; 135° W. long.]; High ridge of banded
appearance 4 mi NE of Braeburn road house; from interbedded coral
reef; Collector: E. J. Lees, 1930. [probably Formation “F”, Lewes
River Gp.; see note under loc. 31, below; Stikine terrane]

Locality 30 (GSC loc. 23418).—Canada, Yukon Territory, Lake
Laberge area. East side of Lake Laberge (Loc. 11 of G. S.C. Bull.
43) [61°15'25” N. lat.; 135°12'9” W. long.]; Formation “F’’, Lewes
River group; Collector: E. T. Tozer, 1953. [See note under loc. 31,
below; Stikine terrane]

Locality 31 (GSC loc. 23462).—Canada, Yukon Territory, Lake
Laberge area; East side of Lake Laberge (Loc. 10 of G. S.C. Bull.
43) [61°14'55” N. lat., 135°11'26” N. long.]; Formation ““F”, Lewes
River group; Collector: E. T. Tozer, 1953. [Tozer (1958, p. 19) as-
signs this and other localities in Formation “‘F” of the Lewes River
group in the Lake Laberge area to “late Norian (mid-Upper Triassic)
[sic]” and cites the occurrence there of several species of pelecypods
in association with Spondylospira lewesensis (Lees, 1934). This age
call is apparently based on general similarity of the fauna to an
assemblage “widely distributed in southern Yukon and in British
Columbia west of the Rocky Mountain Trench.” He specifically cites
similarities to the fauna of the Tyaughton group of southern British
Columbia, which occurs on the west coast of Vancouver Island in
“beds overlying shales with Monotis subcircularis.” Silberling (writ-
ten commun., 1989) assigns this locality to zone UNo2 of Tozer
(1980), based on associated fauna; Stikine terrane]

Locality 32 (LSJU loc. 720-C).—Nevada, Lander Co., [Cain
Mountain 15’ quad.; Augusta Mountains] “W. slope of S. end of

94 BULLETIN 337

Lone Peak [Cane or Boundary Peak of 40th Parallel Survey], immed.
beyond the divide of the first deep canyon S. of Jenkins Ranch; beds
lie conformably on Carboniferous? limestone [= Upper Triassic Cane
Spring Formation]; fossils from top of 500’ thick zone, beginning
60’ above ?Carb. [Upper Triassic] Ist.’’; Collector: S. W. Muller,
September, 1928. [Osobb Formation; zones LNol-LNo3 of Tozer
(1980), based on stratigraphic position; Triassic cover on the Gol-
conda Allochthon]

Locality 33 (LSJU loc. 724).—Nevada, Mineral Co., Hawthorne
[1°] quad., Pilot Mtns.; Dunlap Canyon, 3 mi up from the mouth of
the canyon on the W. side of the canyon; Collector: S. Muller, April,
1928; September, 1934; September, 1935. [lower member of Luning
Fm.; zone LNol of Tozer (1980), based on fauna at this stratigraphic
level; Paradise terrane]

Locality 34 (LSJU loc. 764).—Nevada, Mineral Co., Hawthorne
[1°] quad.; Pilot Mtns.; E. of Mina, abt. 1 mi S. of the mouth of
Dunlap Canyon, abt. 1750’ due S. of a rhyolite knob; impure lime-
stones faulted against slates; Collector: S. Muller, May, 1932. Upper
member of Luning Formation. [zone LNo3 of Tozer (1980), based
on stratigraphic position; Paradise terrane]

Locality 35 (LSJU loc. 800-B).— Nevada, Nye Co., [Ione 15’ quad.,
T. 12 N., R. 39 E. (unsurveyed)]; about 1 mi from the mouth of
Union Canyon, E. side; Collector: S. Muller, September, 1930. Lun-
ing Formation. [zone LNo3 of Tozer (1980), based on associated
fauna; Paradise terrane]

Locality 36 (LSJU loc. 844).—Nevada, Mineral Co., Hawthorne
and Tonopah [1°] quads.; Pilot Mtns., Dunlap Canyon; lower lime-
stone near coral bed; Collector: S. Muller, September 12, 1934. Lun-
ing Formation. [zone LNo1 of Tozer (1980), based on fauna at this
stratigraphic level; Paradise terrane]

Locality 37 (LSJU loc. 872).—Nevada, Mineral Co., Hawthorne
and Tonopah [1°] quads.; Pilot Mtns., between Cinnabar and Dunlap
Canyons at the boundary of the two quads.; low hills between the
two canyons N. of the prominent, andesite, rhyolitic hill; Collector:
S. Muller, September, 1934. [lower Luning Formation; zones LNol-
LNo2 of Tozer (1980), based on fauna at this stratigraphic level;
Paradise terrane]

Locality 38 (LSJU loc. 1014).—Nevada, Mineral Co., Tonopah
[1°] quad., Pilot Mtns.; Cinnabar Canyon, near mouth of first trib-
utary from the east, above the “Slate Gorge”’; Collector: S. Muller,
1936. Luning Formation. [zones LNol—-MNo2 of Tozer (1980), based
on stratigraphic position; Paradise terrane]

Locality 39 (LSJU loc. 1377).—Nevada, Pershing Co., “Lovelock
[1°] quad. [Buffalo Mountain 15’ quad.]; abt. 16 mi E. of Lovelock,
abt. | mi SW of Nevada Quicksilver Mine; brownish thinly lami-
nated shales and gray limestones”; Collectors: Muller, Bush, John-
ston, [E. T.] Schenk and [H. E.] Wheeler, June-July, 1930. [same as
loc. 18, formation not given, but = uppermost Hollywood Fm. of
Oldow, Bartel, and Gelber (1990); zone LNo3 of Tozer (1980), based
on associated fauna; Jungo terrane]

Locality 40 (LSJU loc. 1517).—Nevada, Mineral Co., Hawthorne
and Tonopah [1°] quads.; Pilot Mtns., Cinnabar Gulch, abt. 1'2 mi
above the Slate and Cg. Gorge, last exposure of Upper Triassic
| stone before getting back into Slate and Congl.; Collector: S. W.

934. Luning Formation. [zone LNo3 of Tozer (1980), based
ic position; Paradise terrane]

LSJU loc. 2336-A).—Nevada, Lander Co., Sonoma

juad. [Cain Mountain 15’ quad.]; Augusta Mtns., 2!2 mi

Jenkins Ranch, T. 25 N., R. 39 E., 400-500’ above base of

unit; Collector: S. W. Muller, date unknown. [Osobb Formation.
zones LNo2-LNo3 of Tozer (1980), based on stratigraphic position;
Tniassic cover on the Golconda Allochthon]

Locality 42 (LSJU loc. 2348).—Nevada, Pershing Co., Sonoma
Range [1°] quad. [Rose Creek 15’ quad.]; East Range, N. end, at the
head of Rose Ck., elev. 100’ above base of limestone; Collector:
S. W. Muller, August, 1940. [Dun Glen Formation; zone LNo3 of
Tozer (1980), based on associated fauna; Triassic cover on the Gol-
conda Allochthon]

Locality 43 (LSJU loc. 2546).—Nevada, Pershing Co., Sonoma
Range [1°] quad. [Cain Mountain 15’ quad.]; Augusta Mtns., 2'2 mi
SE of Jenkins Ranch, abt. 400-500’ above base of the unit; Collector:
S. W. Muller, date unknown. ?Winnemucca Formation [on strike
with loc. 41; zones LNo2—LNo3 of Tozer (1980), based on strati-
graphic position; Triassic cover on the Golconda Allochthon]

Locality 44 (LSJU loc. 2970).—California, Shasta Co., Big Bend
15’ quad.; [NEY4sNE“%SW'4, sec. 25, T. 38 N., R. 1 W.; Alder Creek
Trail, abt. 2 mi N. of confluence with Devil’s Canyon] Collector:
A. F. Sanborn, 1950. Modin Formation, Devil’s Canyon Member.
[zones UNol-UNo3 of Tozer (1980), based on associated fauna;
Eastern Klamath terrane]

Locality 45 (LSJU loc. 3148).— Nevada, Churchill Co., [Shoshone
Meadows 15’ quad.;] N. end of Clan Alpine Range, Shoshone Ck.,
Ys mi N. of the road at a point 1.8 mi down the creek from Shoshone
Springs; coral bed abt. 200’ below the limestone ledge; Collectors:
[E.] Blackwelder, Sahni, [N. J.] Silberling, August 8, 1952. [formation
not given; zone MNo2?2 of Tozer (1980), based on associated fauna;
Jungo terrane]

Locality 47 (Field No. HC341).— Oregon, Wallowa Co., Jim Creek
Butte [15’] quad.; NE cor., sec. | [unsurveyed], T. 5 N., R. 47 E.;
45°56'38" N., 116°52'55” W.; crest of ridge on N. side of Coon Creek;
limestone lens in Seven Devils volcanics; Collector: G. C. Simmons,
1974. [Martin Bridge Formation; zones LNol—-MNo2? of Tozer (1980),
based on associated fauna; Wallowa terrane]

Locality 48 (Field No. 56-133).—Oregon, [Crook Co.,] Suplee area,
NENE', sec. 2, T. 18 S., R. 25 E.; isolated limestone knob 750'
WNW of road cut 2700’ N. of Camp Ck. crossing; limestone con-
glomerate stratigraphically below point estimated 1800’ above base
of Brisbois Formation; Collectors: S. W. Muller, [W. R.] Dickinson,
{L. H.] Vigrass, 1956. Brisbois Formation. [zones UC1—UC3 of Toz-
er (1980), based on stratigraphic position; Izee terrane]

Locality 49 (Field No. 56-135).—Oregon, [Crook Co.,] Suplee area;
NW cor., NE", sec. 1, T. 18 S., R. 25 E.; limestone outcrop 1000’
E. of Camp Creek on summit of short arcuate E/W ridge; prob. same
horizon as 56-133 [loc. 48]; Collectors: [S. W.] Muller, [W. R.] Dick-
inson, [L. W.] Vigrass, 1956. Brisbois Formation. [zones UC1-UC3
of Tozer (1980), based on stratigraphic position; Izee terrane]

Locality 51 (Field No. 57-417).—Oregon, [Crook Co.,] Suplee area;
W boundary, SE’4sNE',, sec. 23, T. 17 S., R. 25 E., 100’ SE of ““Pau-
lina telephone” located at junction of Paulina—Suplee road with road
to A. Bernard Ranch house; about 700’ above base of unit; Collector:
L. W. Vigrass, 1957. Brisbois Formation. [zones UC1-UC3 of Tozer
(1980), based on stratigraphic position; Izee terrane]

Locality 53 (Field No. 77-S-111).—Alaska, South Central, Alaska
Range, Upper Chulitna district, Healy (A-6) quad.; limestone and
pillow basalt unit on crest of secondary spur on SW side of valley
of the W. fork of the Chulitna River, 3.80 km S. 89° E. from VABM
Joyce (6363); Collector: N. J. Silberling, 1977. [zone UC3 of Tozer
(1980), based on associated fauna; Chulitna terrane]

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LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER

PLATES

ss)

100

Figure

BULLETIN 337

EXPLANATION OF PLATE 9

1-9. Dagyspirifer fascicostata, new genus and new species ....-..- 2-21.22 eee ee eee tenets

Ny 22

3-9.

Paratypes, ventral valves, posteroventrolateral views, <1, showing size and form of simulated partial ontogenetic series;
(1) USNM 450274, and (2) USNM 450275, both from loc. 23.

Holotype, ventral valve, (3) anterior, (4) posterior, (5) ventral, (6) dorsal (interior), (7) lateral, and (8) anterior (interior)
views, x 1, and (9) anterior (interior) view, x 3, showing fascicostate ornament, globose shell form, small, ndged interarea,
spondylospirid spondylium and paired, apical pedicle foramena; USNM 450276, from loc. 23.

10-38. Spondylospira lewesensis (Lees, 1934) ......- 2.2.2.2 ener e cee eee eee tener e es tener ese reese ee ves ees sees:

10-14.

15-19.

Di.

38.

Figured specimens, articulated valves, posteroventrolateral views, <1, showing size and form of simulated partial on-
togenetic series; (10) CASG 66258.01, (11) CASG 66258.02, (12) CASG 66258.03, and (13) CASG 66258.04, all from
loc. 45, and (14) CASG 66253.01, from loc. 39.

Lectotype, partially decorticated articulated valves, (15) lateral, (16) dorsal, (17) ventral, (18) posterior, and (19) anterior
views, x1, showing form, outline, and reflection of subdued internal ornament; GSC 9619, from loc. 30 [also figured as
Cyrtina lewesensis Lees, 1934, p. 35, pl. 1, fig. 14]

. Paralectotype, partially decorticated articulated valves, posterior view, x2, showing reflection of internal surface of ridged

interarea, and impression of mesial junction of cooperculum lateral plates; GSC 9619a, from loc. 30 [also figured as
Cyrtina lewesensis Lees, 1934, p. 35, pl. 1, figs. 15, 16].

Chorotype, partially decorticated articulated valves, posterior view, x 1; on dorsal valve showing contrast between strong
ornament of shell surface and subdued ornament on cast of internal surface; on ventral valve showing cooperculum of
two lateral plates, with ridged interarea exposed adcommissurally, and apical pedicle opening; GSC 69373, from loc. 30.

. Figured specimens. 22, 23, ventral valve [paratype of Spondylospira reesidei Cooper, 1942], (22) ventral and (23) dorsal

(interior) views, x2, showing strong costae and asymmetric valve, USNM 103468f, 24, 25, dorsal valve [paratype of
Spondylospira reesidei Cooper, 1942], (24) dorsal and (25) ventral (interior) views, <2, showing denticulate hinge,
characteristic ornament of costae, weaker on fold than on flanks, increasing anteriorly by bifurcation in the fold, and
descending branches of spire, anteriorly jugate, attached to valve floor by calcareous meshwork; USNM 103468b; (26)
ventral valve [holotype of Spondylospira reesidei Cooper, 1942], posterior view, * 2, showing ridged interarea, denticulate
hinge, and slit-like apical pedicle foramena; USNM 103468a, all from loc. 15.

. Figured specimen [chorotype of Spiriferina acrotamboensis Korner, 1937], (27) dorsal, (28) anterior, and (29) lateral

views, x1, showing characteristic form and ornament of a Peruvian specimen; USNM 450285, from loc. 16.

. Figured specimen, decorticated articulated valves, (30) dorsal and (31) anterior views, <1, showing effect on ornament

and outline of numerous costal bifurcations; CASG 66251.01, from loc. 37.

. Figured specimens, partially decorticated articulated valves, (32, 34, 36) dorsal and (33, 35, 37) anterior views, <1,

showing variation in form and ornament in three specimens in a partial simulated ontogenetic series; (32, 33) CASG
66248.01, (34, 35) CASG 66248.02, and (36, 37) USNM 106521 [figured as Spondylospira alia (Hall and Whitfield,
1877) by Cooper (in Shimer and Shrock, 1944)], all from loc. 33.

Figured specimen, dorsal valve and fragment of ventral valve, posteroventrolateral view, <3, showing articulation along
denticulate hingeline; USNM 450286, from loc. 15.

81

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 9

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 10

Figure
1-3.

4-7.

14-24.

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER

EXPLANATION OF PLATE 10

SPONGY LOS PIralEeweseNnsis\ (CES I 934) je eysrss ren raererae tT eee ee eee eis el hein ereitecicle seieieicie mere ste trekern cca enerenersietorerec fers
Figured specimen, ventral valve, (1) ventral, (2) dorsal (interior) and (3) posterior views, <1, showing surface ornament, spon-
dylospirid spondylium, ridged interarea, and denticulate hinge of a multicostate variant; USNM 450287, from loc. 27.
Spiriferal(Spirtserina) aliaitiall’and Whitfield, 1877 <5 <5. 6.2 = esis eke, sere occ) e o) > oe ctatersreiel = ehey lay stois © evs) sre ievev lst w= «ee s/o) sevete) yee steer
Holotype, partial ventral valve, (4) ventral, (5) dorsal, and (6) lateral views, x 1, and (7) dorsolateral view, x 2, showing low beak,
wide delthyrium, shallow valve profile, and lack of spondylospirid spondylium; USNM 12671; collected 1.5 mi south of Dun
Glen Pass, Pah-Ute Range, Nevada, by A. Hague; figured for comparison with PI. 9, figs. 36, 37 [Spondylospira alia (Hall and
Whitfield, 1877) of Cooper (in Shimer and Shrock, 1944) = Spondylospira lewesensis (Lees, 1934)].

WE SPONGY lOSpiraparmata TIGOVET MOSS) se cvs crs ceye MNCL oe heer 505 SHED AH. eS SS NEP AE, Sle SAPS NER Site SEE Seis a settistcfeuecetesa elaushateieassea 13

Holotype, articulated valves, (8) dorsal, (9) ventral, (10) anterior, and (11) lateral views, x 1, showing form, outline, fine ornament

and subdued dorsal fold; (12) posterior and (13) posterolateral views, x 1 (stereo), showing denticulate hinge, high ridged interarea,

narrow delthyrium, and fragmentary tripartite cooperculum of one basal and two lateral plates; CASG 60975.01, from loc. 44.

Pseudospondylospira perplexa new genus and New 'SpeCieS: . 2... . ce ee ee ee eee Sure eee ceesenyn eiete weiey eis eee sie ese ene here

14-18. Holotype, articulated valves, (14) anterior, (15) posterior, (16) dorsal, (17) ventral, and (18) lateral views, x 1, showing
globose profile, narrow interarea, and hooked ventral apex; USNM 450278, from loc. 22.

19, 20. Paratype, dorsal valve, (19) ventral (interior) and (20) dorsal views, 1.5, showing circular outline, narrow denticulate
hinge, and surface of a strongly ornamented variant; USNM 450279, from loc. 22.

21. Paratype, dorsal valve, dorsal view, x 1.5, showing surface of a weakly ornamented variant, USNM 450280, from loc.
DD:

22. Paratype, partial articulated valves, anteroventrolateral view, x2, in ventral valve showing anterior protrusion of dorsal
portion of median septum, and spondylospirid spondylium; in dorsal valve showing hinge sockets, crural bases, and
descending branch attachment ridges on valve floor; USNM 450281, from loc. 22.

23, 24. Paratype, partial ventral valve, (23) posteroventrolateral (interior) view, x 2, and (24) anterodorsolateral (interior) view,
x 3, showing exterior and interior configurations of paired apical pedicle foramena; USNM 450282, from loc. 22.

pe Pseudosponaylospira alt. P. perplexa> New Senus and NEw, SPECIES) pieyeyevsyev-are atest ie ate hete eatee etsterer eae 2 ee ected ovetere aia) stetarat ata

25, 26. Figured specimen, dorsal valve, (25) dorsal and (26) ventral (interior) views, <1, showing straight, angular costae and
short denticulate hinge of a large specimen; USNM 450283, from loc. 28.
27. Figured specimen, partial ventral valve, anterolateral (interior) view, x 2, showing denticulate hinge and spondylospirid
spondylium of a smaller specimen; USNM 450284, from loc. 28.

101

81

83

79

80

Figure

BULLETIN 337

EXPLANATION OF PLATE 11

1215 Spondylospira tricosta\ newispecies) 4. --.aa:2 = 202s ee

2"

3-6.

14, 15.

Figured specimen, articulated valves, (1) dorsal and (2) posterior views, x 1, showing size and form of a small individual;
CASG 66256.01, from loc. 42.

Figured specimens, articulated valves, (3, 5) dorsal and (4, 6) lateral views, x 1, showing size, form, and characteristic
trifurcate costa on dorsal folds of two specimens, one well-preserved, the other largely decorticated; (3, 4) CASG 66255.01,
(5, 6) CASG 66255.02, both from loc. 41.

. Holotype, partially decorticated articulated valves, (7) dorsal view, <1, for scale; (8) dorsal, (9) ventral, (10) posterior,

(11) lateral, and (12) anterior views, = 1.5, showing form, outline, ridged interarea, spondylospirid spondylium, and
characteristic costal trifurcation on dorsal fold; (13) anterior view, * 3, showing finely cancellate surface microornament
and contrast between shell and internal mold surfaces; CASG 66247.01, from loc. 32.

Figured specimen, dorsal valve, (14) ventral (interior) and (15) dorsal views, x 2, showing denticulate hingeline, attachment
ridges for descending branches of spire, and characteristic costal trifurcation on dorsal fold; USNM 450289, from loc.
28.

16-32. Vitimetula parva, new genus and NEW SPECIES... 6 ee jes on en cere ta niche ae sie esis = a cls oe a

16-21.

26-29.

30.

3532.

Holotype, articulated valves, (16) anterior view, x 1, for scale; (17) anterior, (18) posterior, (19) dorsal, (20) ventral, and
(21) lateral views, <3, in narrow, deep, faintly costate ventral valve showing high, narrow, ridged interarea; in low,
rounded caplike dorsal valve showing absence of radial ornament, USNM 450290, from loc. 15.

. Paratype, ventral valve, (22) ventral and (23) posterior views, <3, showing rough, irregularly ridged, laterally-rimmed

interarea with very narrow delthyrium and multiple apical pedicle foramena; USNM 450293, from loc. 15.

. Paratype, dorsal valve, (24) ventral (interior) and (25) dorsal views, x 3, showing weak radial surface ornament, denticulate

hingeline, and anteriorly jugate descending branches of spire joined to valve floor by meshwork; USNM 450296, from
loc. 15.

Paratype, ventral valve, (26) anterior, (27) dorsal (interior), (28) lateral, and (29) posterior views, <3, showing narrow,
non-costate sulcus and weakly costate flanks, high, ridged, laterally-rimmed interarea, with narrow delthyrium, spon-
dylospirid spondylium and denticulate hingeline; USNM 450291, from loc. 15.

Paratype, dorsal valve, ventral view, <3, showing denticulate hingeline, descending branches of spire, and supportive
meshwork; USNM 450298, from loc. 15.

Paratype, dorsal valve, (31) dorsal view, = 3, showing valve outline and absence of radial ornament; (32) ventral (interior)
view, <3 (stereo), showing bowl-like valve form and curvature of descending branches of spire, USNM 450297, from
loc. 15. [mesial ridge is silicified crack filling].

33-50: Zugmayerella americana, NEW SPECIES! <0). =< 2 <2 oe oie oie cosine ering aah ha Sica hss ae Ache ea

38-42.

43-50.

3: (34, 37) from loc. 20; (35, 36) from loc. 28.

Paratype, ventral valve, (38) lateral, (39) dorsal (interior), (40) ventral, (41) posterior, and (42) anterior views, x 2, showing
triangular profile and outline, weakly paucicostate flanks, flat ridged interarea, broad, flat, non-costate mesial sulcus,
spondylospirid spondylium and denticulate hingeline; USNM 450301 [same as Pl. 11, fig. 37], from loc. 20.

Holotype, articulated valves, (43) anterior view, 1, for scale; (44) anterior, (45) posterior, and (46) lateral views, x3,
showing form, outline, and narrower ventral sulcus of small individual: ventral valve, (47) dorsal (interior) and (48)
ventral views, x 3, showing spondylospirid spondylium and denticulate hingeline; dorsal valve, (49) dorsal and (50) ventral
(interior) views, x3, showing denticulate hingeline and paucicostate surface ornament; USNM 450306, from loc. 28.

86

87

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 11

‘MUA iiny” oe “ee
ry i,
4 i |

%
y
a

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 12

Figure

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER

EXPLANATION OF PLATE 12

1=5. Zugmayerella.americana, NEW SPECIES) 2... ee he ees oie ene eo mince ts seine et te ae aisle le wae elie) e 0104 oo ve ee eieicieit ie eiieisie

25-33.

34.

Me
2-5.

6-8.

Paratype, dorsal valve, dorsal view, x 2, showing valve outline and flattened median fold, USNM 450311, from loc. 28.
Paratypes, dorsal valves, (2, 4) dorsal and (3, 5) ventral (interior) views, x 2, showing faint incipient costae on flattened
dorsal fold, denticulate hingeline, hinge sockets, median septum and ridges marking point of attachment of descending
branches of spire to valve floor; (2, 3) USNM 450313, from loc. 28; (4, 5) USNM 450302, from loc. 20.

. Phenacozugmayerella mimuncinata, new genus and new species .... 2... ee eee eee

Paratypes, partially decorticated articulated valves, x 1, showing size, form, and outline of a partial simulated ontogenetic
series; (6) CASG 66249.01 and (7) CASG 66249.02, from loc. 34; (8) CASG 66251.02, from loc. 37.

2. Paratype, partially decorticated articulated valves, (9) posterior, (10) dorsal, (11) ventral, and (12) lateral views, x1,

showing denticulate hingeline, ridged interarea, and pauciplicate ornament; CASG 66251.02 [same as PI. 12, fig. 8], from
loc237:

. Paratype, partially decorticated articulated valves, (13) anterior, (14) posterior, (15) dorsal, (16) ventral, and (17) lateral

views, <1, showing pauciplicate ornament, somewhat globose form and ridged interarea; CASG 66250.01, from loc. 36.

. Paratype, partially decorticated articulated valves, (18) dorsal and (19) lateral views, x1, showing form and outline;

CASG 66248.03, from loc. 33.

. Holotype, partially decorticated articulated valves, (20) anteroventral view, 1, for scale, (21) anteroventral view, x3,

showing finely capillate-cancellate surface micro-ornament in ventral mesial sulcus, CASG 66250.02, from loc. 36.

. Paratype, fragment of articulated valves, anterolateral view, x 1, showing relation of mesial to lateral plications in a large

individual; CASG 66248.04, from loc. 33.

. Paratype, partially decorticated articulated valves, anterior view, x2, showing incipient secondary plication in mesial

sulcus; CASG 66252.01, from loc. 38.

. Paratype, decorticated articulated valves, posterior view, x2 [photographed under glycerin], showing spondylospirid

spondylium and posteriorly-produced ventral median septum; CASG 66248.05, from loc. 33.

LZ UGINGY TELA TASPECIES Atartsrerarsescre) crete cnet tence ieted een eey ee ohatoone ote earn cna r= See este case cht a ev suenereLelepens} side seis chats (erehavat= lafolegeKedelorote feels

25-29.

30-33.

Figured specimen, ventral valve, (25) posterior view, x1, for scale, (26) posterior, (27) anterior, (28) lateral, and (29)
ventral views, x 1.5, showing laterally rimmed ridged interarea, globose form and very faint lateral plications, USNM
450314, from loc. 12.

Figured specimen, partial ventral valve, (30) posterior view, x1, for scale, (31) posterior, (32) anteroventrolateral and
(33) dorsal (interior) views, x 1.5, showing spondylospirid spondylium, very weak lateral plications, and globose form;
USNM 450315, from loc. 3.

PANG CCL IG ISS EM nog na pba bic cocopdncd oom Gtioanracds pAdonbdnc HouULSe ro OsuonoSoaonOnonee CAndooun nooo Dy abpuDaDo dT Oc
Figured specimen, partial dorsal valve, ventral (interior) view, x 2 [stereo], showing mesially recurved, anteriorly jugate descending
branches of spire attached to valve floor at midlength, USNM 450317, from loc. 1.

103

90

89

89

104 BULLETIN 337

INDEX

Note: Page numbers are in light face; plate numbers are in bold face type; the page numbers on which principal discussions occur are in italics.

@yaldefined|| \seess soe eeee ssc ose tos eee otras anes salen ears 74
acrotamboensis,
SDINULCHUM Cl metas ere a ae eee e ee aN ace 81,99
SpOndylOspirae ee ee ELI 83,84
AERO GA ire cot onece con econce: Gate tnns Snemstendousectecewde nose rasteuenet es 68
A BORSA SAVE ws te di aace va soe aw ae ae salsa a wate saaosndseeasewaslvewsde cities sa sacaa'es 63
Alaskan sees sin eens sci d.72 se eee 68,78-82,87,88,91
south-central,
Alaska; Ranges. ance erncseth- cosas tenet eee tne ans Saes sess Steenes 74
Upper Chulitna district, Chulitna River ..................... 94
INabesnaliquadran gle; recesccsescnsescnss-cacese -a2scse-cneeeateceeeee 733
INutzotinelountalnsipecs.csceeces se seccerseceeeece ec eeceen on eceeeeeee 92
J BSE AG) al) a Se P epee CEE ep ERB ERE ES SHOaEE PEC CaF eR Ce Aue ne ceaede se 92
Copper River Region ..... OD
NACKIG LEEK Eee wes ences cer ecesceceres cdnenncs oncescunes Sotvenertcs too 92
SOltheastemnrss eee eee eee eee ere 68,73,92
GComnwallisfheninsulayec.cssessesce terre eee ease ee eer ee eee 73,93
Gravinarisland! yo: tics coos se ten seeme oe oat oem a cs ysensheweers 73
eekuiislan dl Pr vct cece cassenrseess anceeeeetissedsctesachecactssscnecers 73
IekutS traits | ooic. codec soscesencs cssecetes soe teect sotseesctsrscteuveuve 92,93
INGKUSISI Ot SBeernscee ssucs econo ee ars sesaccnecsetees er enscercscaete 93
thiangulationistationiGORIN) wicsecesceeescerece-secscecreecueeres 93
triangulation station LOW 52,9 93
tang wlation’station MEWIMUeicccresc-.-c-cceccecesscoseccers 93
Kine Slan ds eens. eee es reece eee rane eects crc mee 73,92,93
Pups sland yrete. te sscteecccsesce cots centerenenc saccecsecncvcwscrsescesees 92
ScreenylslandSyescrcnes cnceces crores sere eee ease nee 73,92
Glarence|StraitEtolinul slandiseeccs.ssei cee sete eas see seas eaee tee 92
southern,
Wrangell Mountains
East Fork Valley .......
Green Butte Ridge ....
Alexandenterrame sn. .-ctesccancsresneneteseeeee ss
Admiral tysubterrane tecccec. esse vec teerteree seer core ee eee
Annettesubterrane mete fe see atone ee tec ere ee ree eee
Graipisubtenane pessssscccicace sc sence cota caiorecetes stueeednoceeie sre tee
alia,
Spirifera (Spiriferina) .... 81,100
SSDONGYIOSDING....00.+.500500-< ... 64,81,83,99,100
ILA (alta) eS DONAVIOSDIV are recone eee terete meron seo eee 81
LDS qaeere cerca eee en een ea Tee Cnn oRe TE SHOR eee eo raese rate roee eres 68,87
americana, Zugmayerella ... 11,12 ...... 67,86,87,88,89,101,102
Antelope; Sprines#ROnmMatlOn!:..cscscseecesencastarsatteceer ese access 70,93
Aramachay, ROrmationy sn sctes-c.scevesosseenesenetiods cotenatee reoneerene: 74
Auldiicang:Syne!Groupieseesssesees soaks eee enero reece 69-71
Dun Glen Formation ... 71,85,94
Osobbiormatione.. aamcccesecessoteneeeeens 71,80,82,84,85,91,94
WANNneMuUuCccasEOLMAatlOnimrcssseersee teens eee 71,85,94
Austria; PisGhenwelsersesw esc. ase see aeeesee ae acenae aero eeaae 93
fustr hyceras austriacum Zone [Zone LC2 of Tozer (1980)] ...
se eee decasvecsescnseteeee erate ers COOLS 2199

and Speed (1990)
Baikans
3egg Formation
serg, Jones, and Richter (1972)
Big Bend 15’ quadrangle
Bittner (1884)
Bittner (1892)
Bittner (1902)

Blackwelder; Es <sccccccouuos ss0esseescsecseese <ovecve does eee decane 94
Bollibokka Mountain 15’ quadrangle ..................0..0.ceeeeeeees
Boreal région. 2-305. Jeccss. oosscvesentesssesceeas dees ogee ose ee
Braeburn! Formation 2 cc02i2.502 52. esos vs tede vec ee coke eee
BrisboisyROrmation! sa-sastecceses ees ceeece se eon tee
Brock: Shale: s2.<<.2.c22 Cosserscnecse st acs esse ooce ene se eee eeen
Brueckner and Snyder (1985) ............-......+-
Brueckner, Snyder, and Boudreau (1987)
Buddington, A. Fi 2¢...35.c:cesccees cesses acc2sceess.cose ce ee eee 92
Buffalo Mountain 1'5’quadrangle: .......-........--.-seesseeseeesere 93,94
Burke! andiSilberling ((1973)!\-...<.c0.-.-cccs+ece-ceees ose eeee eee 68,71
Burney 30’ quadrangle’ «.. ...:....<.<cssc0.<ove0e=c-n cess eee ee 92
131)» eae anes or nee re Pera eres scutacoctseccaccnecececo ac 94
Bychkov and Dagys (1984) ..... .. 68,81
Bychkovretals (19576)) 220. ees cne- eee eae oc ies eee 68
Cache ‘Creek terrane: 5.2. ...2.00-2-csccceceesesdetcuse tee nte eet ee eee 69
@ain Mountain) 15’ quadrangle’. -c-...c0s.-«-eseoceeeeeee eee 93,94
California: (co soc nc ecccssaesassonssee Mee eee 68,81,84,87-89
NOMEN ss fs sces ssc sever Doseavede nese sacetsenesesecen eee eee 71
SHasta''Co: ccsdesccsteiwes oe cess cavecevcowneueseceeae ons eeeeneeeReeRe nee 71,92
Alder: Créek Trail) ..52.ceese2s <cccaenns acess cance treo ee eee 94
Beari Gulch. =. 2scc2 seen occa Ser coc caecudes st ee ee 92
Cedar. Greek: ec s.c0 02 cccess sates cob ecsnun see eec ste 92
Devil’s Canyon .... 94
Furnaceville ........ ae 192
Grizzly Reake-- ssc... sey 9
Little:Gedar Creek 2.225, 2s: ccccacc sect see 92
Little:\Cow Greek. s.2s.ccuscscs. Tess sadheseecesee eee eee 92
Round) Mountain) ...05.....6. oce8 sacakeed coccos oe ee eee eee 92
Campbell (1990) c.¢ .cone.dec eG. eee boos oeeseeac ee cc nee ee 68
Canada,
British Columbia,
Halfway Riveriarea .....:.....<cs.ccosece seseeece ee eee 66
NiGolait sic fisccoeeecaws Laaecioac a eSoesonc Sasso ee ee 65
southermsc-teeseeee see 93
Vancouver Island ... ws 193}
@anadian: Arctic, Cameron Island) <2v.c2s. eeceseseceoeeeeeeee eee 66
Yukon Territory,
Lake: Laberge repionivess. -ccore-ce-paeeteeee ena steee eee 66,72,82,83
Braeburn’ road house .........<..::c«chesecssssseneesasseaeeeeeseee 93
east side (ofillake Laberge): .-s.cc-c0. ---eesseeeen ee eee 93
Ieime/Peaketerntocccsenec cee ..- 66
southern ....... 3 293}
MroutlLake y-cc.cesacsesee eae vee x (66
canavarica robusia, Spiriferind .icccc.csccse.ce.cceeaseseevsteeteeteeeee 81
Garpathian Mountains) <5. 2seeeecenecescersecsces costes toereaereeemeeres 68,87
CASG [California Academy of Sciences, Geology Collections, San
Branciscos\ GA) ssseeeeetee eee 74,82-85,90-92,99-102
Cathoart;(S. Hi... .cowccssooses sree steer ote 58 S02 hee Oe 92
Chinavree.<soecacesoune ... 68,70,87
Get Zhow «; 3.2. caccececs sods adacsac se sos aeseeaase soe eae eee 67,89
Ghingyand! Reng (U977)) ee ccceccsneasese eene eases a eenee Renner 68,89
ChinleyBormationie.eese teres ee vans OOS
Chitistone Limestone Formation ..... ... 73,88,93
Chong’ D: and! Hillebrandti(l985))e-..cccececocsesseeeoseeserecee tear 68
Choristoceras crickmayi Zone [Zone UNo3 of Tozer (1980)] ........

Juadneiee vededacdeasccseceesk cevcvesewesteaseneoee 66,68,81,82,84,86.92-94

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 105

Chorolypel|Gefined | pesraccescesesccecccccetccescceaccscce cc ocesscccecnecones 75
(Ghulitnattenmane ress... tocseecsec crores ctecrsecstessscerccress 69,74,82,94
Glovem Greeks GreenstOneiecsssster ccae rece netesecececeearcesmettcnee: 72
Cochloceras amoenum Zone [Zone UNo? of Tozer (1980)] ..........
BRN aC cece eas dwone nse cagsucevesenessbesusactstsete 66,81,82,84,86,92-94
OETA EOC Goer eae coe cee tenur ras acdesten casa secacc soc cbeuncumaceresne rates 63
(oloradoyblateaw region sc.scecsecseece cece scnceseceecatcceseececaseonseasrs 71
columbianus (cf.), Tropiceltites scons}
(Crayss yO (DIKE) cneanascedgensceoancadsessndesna poche sascaebacaseceadsonenoac 71
Gondorsinga MOLMAatIONGstesercescecccceres cee scce-cececenec-neeccetanases 74
Cooper (1942) ............ 63,64,66-68,74,75,79-83,86,87,89,92,99
Woopen(MOA ayo cciccsiccscacteccenctvecev-costubaccscctes 64,81,86,99,100
Gaopercuiumililustrated|| recesses -c e-cceecarecsotecsceccsssssosercceee sm 76
Horde vse al MOST) ssccccsee so re- ccs ciacseassork secs se Mesccescsveoneece ses 69
Wordillerawre eet ro ecco e es eo cere canen es soon e gtases devaesucense toc cescs 68,69
WESLOMI ee ene ce te ntice a acee cote ene ee hn canon ee Soeme comet ces cCoteeeetetees 68
Cordilleran
DEV al WeSteeacce ce ccc se sectwdcorsscocesatcn sche ccs cncees Suet uecncectconseresess 69
COTTAN CS eres sere cccccris ster sees cacdsoccct seene steers seusciecceeseoae 68,69,74
(CHTITTGA, « serscnbe SSCS ESSE A SAECO aaa gen 68,87
uldesacelibmquadranple eeccs soccer ree re cree eee ee nec eestenetects 92
GyriinanDaviGdSOnyl'S58ic-c-c.00 coves scec ess oce tees enc cnessteceeceses se 64
LEWESEMSISNECES) O34! esis conten nse rorerecensceateaccentecere 81,99
Cyrtopleurites bicrenatus Zone [included in Zone MNol1 of Tozer
(WOO) | pcssandscsasecnccksasssanceresse neces ceeeaeeace 66,80-82,84,9 1-94
GEOrbigny (S47) Nasa sees tet ocen cr. ates oan ch eceteraactodeucecodeseosees 64
ava [defined] Pree ehos sd sotceehscaces vaeees sce cesecsclateccscuseu scsumecncues 74
MARY STG OD) Hasac sh sonececsnmensceete sean cuectnc or cscuacerearocecoteesss 81,83,87
ID AGYS! (USGS) fe casenen.cs-neenaeiesaerecee-seseees 63,64,66-68,81,85,87,89
DD AGVSE(UOG'S) Ae csetcnesecae secicenodeemaeswanedenedvedeecetes 77,78,81,87,89
DALY SH (OIA) eesete sek ae ode dees esctaseccceseazesssmdese case 67,75,81,87,90
A GY SH (UGH) Recketaat tucesedsconeceseseseqssusaeecuedveneaeteewenedese ceeeee 89,90
DALY SHARES i iocece sete ccweatns de eawacsieuacemeceecttenes 63,67,68,77,81,82,87
IDA SYSDIFIFEr AM. BEM sce cen sue cceaccec-easaseccesetcsceesees 75,77,78,79,90
fascicostata, n. gen. and n. sp. ... 9 ...... 63,66,67,77,78,89,99
WMavIGSONI (USS —1886) is osc scsececosemoeves accesso seeeenceeceeeteeceee se 64,75
Debiche, Cox, and Engebretson (1987) ...............:0c:0eceeeeee eee 74
Department of Geological Sciences, State University of New York,
Binghamtons (Nie cccccsssccesscsesesecesweveveteceeceee se -cee-cessceeeee 63
Dickinsoniand Vigrass|(1965)) -...c.0e.se-:-2-2e-0 $ec20-occs ee oenteeeees 72
Dickinsons.W...R.........:.2ss0+se05s 94
Diller; JecS5 ...:. 92
Dinarides ........ ae wee aa a we 87
DOV eTa 99 (0) Pesce aces dete ese aerete deen ee ten Sateen ee eae aeaas 70
Drepanites rutherfordi Zone [included in Zone MNo1 of Tozer (1980)]
BN BSD, ced ccwceaxiecuceance sen vascesee tees’ 66,80-82,84,91-94
DD tT O Melee eco seca sescecnecsavdece sscdebwateudascestes tes gacedecteeecese 63
PaplexGreekaROnmatiON, <22.csssceseseee-2s0sceeevsescnce-cecteeceeeseccesee
Eastern Klamath terrane
MANY INO SS) iecssesssestencecceocst cnc ccccceeuscuscccsocesh toe esdeecnoreeee
Enterprise 15’ quadrangle ms
EUR CRZ UTINGY CNEL ea soa aise sos ian sn sas at see acids stone awa Ps new nana «ee
Family
Sy TUN d aes seek ace ced occu otacsuwa sede awelane sa seancesseccaceedeome cranes 67
MaballidaeDagys;, 1962) ..0......22..--ccsene--2- 0 63,67,75,77,80,87
Spondylospiridae;n: fam: .<.220.....-.2--.<..sc0cecesasee 63,67,75,87
fascicostata, Dagyspirifer ............. Opec 63,66,67,77,78,89,99
ENCE MAKE Tat GUStenceese se oacesacseceacee eee seat eee oe Bee re eee 70
SRO, 181k (Gr Aaa eae arn er ee oe Ent Bt 92
Field No.
DOa133i(Wocality/'48) mecsescseecetwuwdecsocsenscacaenesacneee 67,72,92,94

HONS 5((Mocalityr49)) <.socanceccs-nsceoee-caeuconcs-neseennn 67,72,92,94

57 =A M(EOCali tye oil) Mtccctcecesssesesseccsessaceseseacsecse. 67,72,92,94

Ti=S= lie Mocalityas3)tovcsscccccscsecccce eaetcececcccesoes ses 74,82,94

HC341 (Locality 47) 72,82,94
RortiethubaralleliSurveyarcessescccsscrceseccteececee ete tee ences eeeeeteses 94
Erizzelli(lOS3)horeccaecoracerettee ccc reere ee ee ter cece toot mncer erence 75
Gabbi(iS64)ierc.cccssccstceceneces see sace seer ec soncts scar steeree cnese scons
Gabbs Formation es

middlesmember Peres sicsc eaccco ses cecaes.cececocecs ance cers sseseececess 81
Gardneriet"ali(1988)) sores crcccecencce soc cvsass sere cocesudsesstensese 73
Gnomohalorites cordilleranus Zone [Zone UNol1 of Tozer (1980)]

SEEPB AL bps 1 ala Breda Bad ee sero hon= 66,68,80-82,84,86,92-94

Golconda Allochthon 67-70,71,80,82,84,85,91,94
Gold Range lithotectonic assemblage of Oldow (1984) .......... 70
GraylocksFormationt .6..5.. ccsscoonccnscnscosce.ccsnssdesererese teeer aces 72
GreatiBasim 22. osc os: cf ccs = ooo eeeee teens ss Goes e eee Sesto eee eee ences 71

GSC [National Type Collection of Fossil Invertebrates and Plants,
Geological Survey of Canada, Ottawa, Ontario, CANADA] ...

PEN het hers Agoe MON a ote eNee ea aed Seis NAeTES 65,66,74,76,82,83,99
loc*WO229i (Wocality:29)i\ fern. -co-sseecesesceeseeseeece ese tees: 72,82,93
locy234i8i(ocalitys30)ieeters-c-s cores cece: 72,76,82,93,99
loce 23462" (locality 73il)) peseseesese see eeetee cece eee 72,82,90,93

GSC [Vancouver, British Columbia, CANADA] .................. 63
Guembelites jandianus Zone [included in Zone LNo1 of Tozer (1980)]

ee etd honey SRS. 2 sane a cate 66,67,78-82,84,88,89,9 1-94
GuextandiMaylori(lOiG) i. ceccsscoscseosctcccnctecrteercececeeseetenete ces 70
Faas? (OSB) i ease eccceccec oacac ncn enee cca sees re hee eer ee Te eee 72,92
1g F120) Nike ener Re eRe Oe oS EER Onc OCs EE RIac Ona ceaconO la naeondnoconacqcnch 100
Hall and Whitfield (1877) 64,81,83,99,100

Hallam (1981)
HH Globiaieeccsecssesnececassse
Halorella Bittner, 1884 na Ae 5 patieaas
HarliButte IS aquadrangle secs -ceccsdencce secrete caters rete
Hawthorn’ quadrangles ee ese se sccsenecesce eee eter cere ceee

Healy A-6 quadrangle
Hector (1879) ase pe
HCI €S os cesccsicsrescscesecssscescuecssses
hemipyramida, Zugmayerella
ETIHOUSE NOT Te ccc cote cceccse wactence cect ectercesaceat cere cc ssecctrcccwe
Hillhouse and Grommeé (1980)
Hillhouse and Grommé (1984)

Himavatites columbianus Zone [Zone MNo2 of Tozer (1980)] ......
BEBE EBA S8 cE IIR EH EERE REE BECACaRn ERE ACiry BicCEnoa Bo 66,67,80-82,84,9 1-94
Hollywoodsi ormationyess estes. sents cee cee cer scree sce cesonecsec ses 70,94
Homestead 1:62!500! quadrangle’ 2vic.ic..-ssceceee-seaeeeoceeeee cece 93
Hoover (98S) eeeeerre rer ereoreeee ee 65,66,68,79,80,81,83-85,100
13 KoYONKS ef (VICIS10)) ae reer peepee Cec neonncadooreacEch doce sea ar oaarcccod 65-67,74
PLOVER (199M) Pe eccc strc cee cceerasean crass ivseceasenctetcesecess 77,79,83
FIGOVET! PR ee eee cate a eanenctececctcc oreten eset erenaetetereee 68
Hosselkus Limestone Formation ........................... 71,88,89,92
Houseiand Sentor (1980) cssesccstececceeteen ene ene ceecen reece eter reee 66
BlurwaliRormiationic. cre cccscscctecnce ea ceea tes eee cere ene ee ene eee 72
ELV GiGroup ies rece note ecco se tec cece tee estat acceso rr eer ce eee eee 73
Burntilsland(Gonglomerate mess. sore ssee cere ce se ecee cae neeee cece 73
Cormwallisdamestone ee. -essee see see eee eee eee 73,78-80,82,92,93
Hamiltonvlsiandslimestone ic. seretes seco ce ncacseesscenceceemescces 73
Hound Island’ Volcanics\re-tce-:. sencsccsesesesescctaceecccesece 73,80,93

| (oF toe codensaE ence cobantaacnoce Rone ane tier er ecrasnanbacc 68,69,81,83,86
WWE WIS COs eae ce Si cheat te ee 12592
INGZSRELCEKEO tyrer sees eseee ee sere tee eects wes 2-92
JaqueseRe RA Station. coseacsecosscesnterseaeee sass saneee ce eesteneses 92

Mi ssioni Greek 8, oi conse sideccuaceacde sna encesseonsean@etadeccetavases 92
Slickpoo! Settlement o.-..cc2ce.c.c-cscecvcteosencnssecsctececsceecess 92

106 BULLETIN 337

Idaho, Nez Perce Co.

Staoseph?s Mission 2.2.0 .se-eesesesee ace one toe acesincmee Saoseecsaeses 92
Nez Perce Reservation east of Lewiston ................::0002008 2:
INGEQUIDIICALA MZ UCINIAVCLEll Gi rerrene ce tetra eee ear ee eta 89
incinata, Zugmayerella 90
Institute of Ecology, Vilnius, LITHUANIA .......................2. 63

ToneuliSwiquadrangle) sete scccc sec ccncsce toons -eseececccnsaniete eee seeseseeeees
Irving and Monger (1987)
Inwinhs Wess ees:

RFECKIENEANG ben ceescsee eee ceatcceerccma-ccectecean asecitecmen
JimuiGreekoButte lo vquadran gles. .c2.0--s-ereseeecretes scene ee see nen cre 94
JOHOSONM Raph tee eace nen cctoeass accecscenencetere sactaseceraececeteresssecss 93
Johnstomiersces ese escsrs se rencesestore

Jones et al. (1987)
Jones et al. (1980)
Jones; Silberling, and! Hillhouse (11977) -.......2.2.-.-.-2.-2--2--0----. 74
JUN GOIeMANe seeseesceec cece cere ce eee ceene == 67-69,70,71,79,82,92-94
Juvavites magnus Zone [Zone LNo3 of Tozer (1980)] ................-

Bb cS SCOURS EEE oO CSE TEES 66,67,73,80-82,84,85,91-94

Keku Volcanics

Ketner (1984)
Ketner (1990)
Rain Ge SEM 5 oo arc Sa yeaa oe ae anaes» oot ee eee rae ae eee 92
Klamathites macrolobatus Zone [Zone UC3 of Tozer (1980)] .......
Be reeset eae eee 66,67, 80-82,88,89,9 1-94
IRiipsternellanDagyss 191 der eccsrcntanteces seer eeasestes--casceeeee sate 75,87
koessenensis,
SS DIM ISCTLMG tes ceaeen erated eno atets Sea see catou vot eascewecsseecssccssecseers
Zugmayerella
Koren (19317) eres secc sess eeeee
LaballasMoisseiev in Dagys; 1962) ....-.:c00<+.0-cn0+--0-<-2s <0 81,83,87
Laberge Group 73
DeaAWSi (982) wetorecdocns ccnees ace cdecysseeee cstwame cxogecnseaaoees anes testnasse 70
Mees (1934) \c.cc.ccesee5se 64,66,69,72-74,76,79,8 1-83,86,93,99, 100
DOS NIES aa ccstsosscs ies ssoe nemo notaa seonshteaceavsesis dhe stdvddesceaueedsasedsce 93

TEPISTNAliNG WANG 19S Sg recscendeacotes-accseaesceenecnceevesseavecceceses
Levy and Christie-Blick (1989) ...
Lewes River Group
Formation “A”
Formation “B”
Formation “C”
Formation “D”
Formation “E”
Formation “F”
Formation “G”

lewesensis,

GYNLIN Gig ceaneck seo deus vaceecsescewuetasSeacwe nce ncdet cu ceasesmacewess 81,99
SDONAYOSDINAS Fcc cce-u¥-oa22ae 50 4scouses scosuceicccn cence see den suteswimsemanses
Pe ct: 910 ...... 63,64,66,69,72,74,76,79,81,82-86,93,99, 100
lewesensisi(Cf.) sSPONAYVIOSPINA. vo... <sc0.s-caecceassveectee-snaescce=— on 81
ee wis: Rog Wiser: Se Peis eee as cae get es ete eras ena ea 93
Rosttles (1 9857)) coccsccce eacec cc conc eee co scee saneeens coun awe aces aaestaeeanamenens 71

Localities used in this study
KS )106: 2309) ieee corres eee eae 71,89,92,102
ic 71,84,92
GS loc 71,88,89,92,101,102

(USGS loc

71,81,92

6 (ISGSMOCs 1054.8) ire see ceeencseceaceeeeee ene seceee 67,73,81,91,92
MCOSGSocsW406)\ orn srs, csceceaconassernceeseese nese ener TST O292,
Si(USGS oes S14 ee wecccwcccsecerecenseccesccaeseoees 70,84,85,92
10 (USGS)loc! 18354) neces ceoeesececesce sect cere ee ee 70,81,92
12 (USGS loc. 15550) .-- 70,72,89,92,102
13 (USGS loc. 16266) ............ Se deasiswsen ton tert eteecene 73,81,88,92

15 (USGS loc. 17424) = “the Lewiston locality” ....................
See ee ease oa pe one Na ere seen 72,8 1-83,86,87,92,93,99,101

16) (USGSi1065243 85) essences sacs ceeee sae nescence ace eaeee 74,82,93,99
EPCOT G SS Mores TN PYG) cecccocacacancedbaassnaccbcansscaccocaces 70,79,93,94
20 (USGS loc. M1708) 73,88,93,101,102
ZAM(WSGSilocy MIN749)) <ec-s.cctesecscne ates cece esecececes 72,82,88,93
22i(WSGSi loch MiG 06) eessecesewesees see tesco noes 73,80,93,100
23 (USGS loc. M1911) .. 73,77,78,80,93,99
DAW SGSTOcaIMUO IW) secccccsoenes conc eeectee eeeeee eee 73,80,93
2'5\ (WSGSiloc: MiG 19) ions. ce ccs se e-ce-encosceeneaensecseaeese 73,80,93
261(WSGSilocs M2135)\ cesccescsses eee wees eee seeeee eee 66,73,82,93
PP WORN GES Ties IN ANEIS))\ caceecsasecsoonceneeccnaccoace 73,80,82,93,100
28 (USGS loc. M2672) 72,80,84,88,93,100-102
29;)(GSGiloc: 10229) <2... =:es2-cce cc costes sce esce 0 eee eee 72,82,93
30\(GSGiloGH234 18) eecccoscecw ccc ste essere 72,76,82,93,99
31 (GSC loc. 23462) osiow sede deeee Seen 72,82,90,93
32) (ESTUplocw720=C)mececse eens seo . 71,84,85,90,91,93,101
33 (LSJU loc. 70,82,91,94,99,102
34} (USTWMOCAAG4) eeeascen.cceencscacrescsseecseteenee teeter 70,90,91,94
35i(ESTWlOC4800SB)) c.cc22cc-cereacee neseeeseer eee 70,82,90,91,94
36 (LSJU loc. ... 70,82,90,91,94,102
317) (ESTE OC! 872) res seseawscesnes ome reer 70,82,91,94,99,102
FSO Coretan (0) 12: PB aneeecerescarecoscopeecubscanoscceoce 70,92,94,102
39ST L377) nxs-s2cc- 2s -seter ence eece ee ee eee 70,82,94,99
40 (LSJU loc. 70,82,84,85,94
41 (LSJU loc. 71,80,82,85,94,101
42(ESIUW 1ocs 2348)) cacvcccccccnocescecsecseeecaematoreees 71,85,94,101
43 (STU 1062546) .cc.s cece hee cscse sees eoc ose ce ere eee 71,85,94
44 (LSJU loc. 71,84,94,100
451 (ESTUPocHsil48) ie eeeceesrecree eee eee 70,82,92,94,99
AT\(Bield\Nos HG34 1), ..siocc--consee sso ceeceseses coeeee eee 72,82,94
48i(Field(NO< 562133) ioiis sececcc case eee aecenccntone tens 67,72,92,94
49 (Field No. 56-135) 67,72,92,94
51 (Field No. 57-417) 67,72,92,94
53\(FieldNo: 772S=Vil)) a2: -cessecssvnscccesse eee eee 74,82,94
Locality 10'of GSC Bull! 43) Sires san. seen reece eree eee 93
Locality 1 liofiGSG@ Bull43\ oo. i ie recace wees soesnesconsees erent 93
Lori Formation .viccicccceo0sceacsocsecen tes estescns poe eee sone aoe 70
Louvieres [48.2 tiad. eee 63
Lovelock 12'quadrangle:< .c......cc...c2sn. cee oer seen ee eee eects 94
“Lovelock assemblage” of Oldow, Bartel, and Gelber (1990) ........

Lower Sedimentary Series [LSS]
“Lower Triassic limestone”’ of Richter (1976)
LSJU [Leland Stanford Junior University, Stanford, CA] ....... 74
LSJU localities
720-C (Locality 32)

71,84,85,90,91,93,101

7124) (locality? 33) + foeettes-cocecesessssecesceencs 70,82,91,94,99,102
164 (Locality 34) Gi. ccttccscetocecssnorces = sacceecaseeeenees 70,90,91,94
S00=Bil(ocality:3'5) cscec-esness--eoeeacteeeacceceeecenes 70,82,90,91,94
844 (Locality 36) 70,82,90,91,94,102
872 (Locality 37) 70,82,91,94,99,102
HOA Meocality/38) ieceice aeeces ee wesc snaes-oo- erect seneenee 70,92,94,102
137 7a MWocality39) seccncs-csccrene sade’ ses easeeesneeeereeee 70,82,94,99
157 (ocality:4O)) 4 acces ees. eecececesteaseseaeesse-e: 70,82,84,85,94
2336-A (Locality 41) 71,80,82,85,94,101
2348) (Locality:42) leas csserccscececers-e-s cee eeeeeeereees 71,85,94,101
2546) (Mocalityi43)\cesccssseeecaecaecweneeosue concen etece meneeee 71,85,94
2970 (Locality 44)
3148 (Locality 45) eS

LSS; [Lower Sedimentary, Series] =-.-<---.-2-c-2s-c0-<eenserenecsesenesene 72

MuningsRonmation\-cessess ese eee ce aeeceecanes 70,82,84,91,92,94
lower members. -222.<cceeaosdencesoesoeree sense rece aces eemeeeee 91,92,94
Upper MeM DEL ...nscc-cas-eecesceceere seen ssaees eee 91,94

Luning lithotectonic assemblage of Oldow (1984) ................- 70

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 107

Muperand'Silberling (985) tecccse-- es ecee see seen nee e oe ete c ee reels 69-71

Malayites dawsoni Zone [included in Zone LNo2 of Tozer (1980)]
EE sea nx seteccet nese tas dean 66,67,80-82,84,85,89,9 1-94

Malayites paulcki Zone [included in Zone LNo2 of Tozer (1980)]
66,67,80-82,84,85,89,9 1-94

Martin Bridge Formation .................... 72,80,82,84,88,89,92-94
Mayzandi Butler (11986)).<c2..2..cs..--+-2-eec<csseceuecessececesecereenonce rs 69
McGarthyskormation as-cosccercee: caceorccercae es -cct recs ceeeeaeaecteteoes 73
IMAG) STV OYES IS 1825 8 a anganscsoa se saaaa sce ascacanscosatedotacosasacacsGsp60I00050 63
megalodontidibivalVesitc..2---ceccceme cree cece se ces sacesoaneancssaeeeseaee 74
IMMillerenaln (M984)! cc.covesseccwacwes ssecduactvc oeneceuses-Seseesncersees 69,71
MitlivallemiSiaquadrangle: i :c..4.ce-aceceesseeeee ce eee sec clvese seesitcesse 92
mimuncinata, Phenacozugmayerella ............0.c0cceceveneceeneeneeneenes
BRNO ron ois nama ase antea peeeere swe 12 ...... 63,67,89,90,91,92,102
IVIOGHIZUKANED WEss boat. cscd sertevs sie ccon ate cstcncubacanowenecususenasenspene tee 63
IModinW@ROrm ation) ca. csccsecse-ne cece nc ieeenaets steteascedeaneeetectenes 81,92
Wevaless€anyvon Member eres scc-cs sec sces-c-e ee cseeeecssctensee 71,84,94
Hawkins G@reekiMember <:2¥.ceccesccastsescesesess secs secsceesecrcesses 71
Iroska Mem Demin seiccccccnwstenccb tess stescscaesscncsovenscecsecctearterces 7A
INTO fri tame el petite sche aca s on Secah cose ceseciee se vescseeetsdur ateeececeeaeues 92
INTOISSCLE Vg (UGG 2) be merci bcos teas sesevene tes coeesenaeesseacarceesese ea 81,83,87
IMO] SISVIGSILOS Wat si race 2 onsets oc sale osw sedate veteseteedesevesesuescuctweseses 93
Mojsisovicsites kerri Zone [included in Zone LNol1 of Tozer (1980)]
PRD ee as chiywaa seus ie seateseserse 66,67, 78-82,84,88,89,9 1-94
Mongermandiberg (198 7). ce ose escent ee ce ee see en ee erecee eee 72,73,92
Monger, Price, and Templeman-Kluit (1982) ....................0.. 69
Monotis subcircularis Gabb, 1864 .................000.00c00eeeeee ee 73,93
INTO OTE WB HIN acta ca cecee ce cheno cc ods cccka ces utoncuouesterconcnescseeses tones 92
IVA erat (A119 G17) eee ee cocoa os ss ocae oo cue wencitne sec ceseneeove ceesenee 73,92,93
IMU er eesti es cece cccuccucscoccesarnosse toonstuoeeaeocaeenene eet cceeemees 93
MullemandtRerguson(1936)) ..s...cc..scenececeeee sere seseccesecceteme ses 70
Mullemand! Ferguson (1939) ....c.ccsccncsssescestecceocsrateceesarcsrsese 70
IMLS TRIS A WWiojs ro oo seeoo e's cote oneawes Seam aivesie neds eeumechomeneteesseeeceeee as 94
Nabesna C-5 1:63,360 quadrangle ....................cseeeeeeeeeeee eens 92
ING VAC AlN sone eer net Seow rakCarcnascbancedenccvens 68—71,79-82,84,85,91,92
Ghur ch Co MR ee. oe csc-wcowe cue caesar ena seascn-useatenaseeseeceeeets 70
Clan Alpine Range

Shoshone:Greek- 7s octet. set Se RR UR kod cote
Shoshone; Springs yecscccnc-c cost 2 eee ees ese eoceoeeoew sae ennee
PATI GET C Ospaneeoe Pecenoias on clave tet cto ee
Augusta Mountains
Boundary Peak aa we sia
Wane Rea ken. oe eek oes sanens suacsgansacstancceveecsdeseecss eeeteesses
MenkinspRAanch i io25 5. sr2-2ds ce cccesnscesecctsescncsesearssseastceeeeves
Lone Peak
Mineral Co.
Cinnabar Canyon
Dunlap Canyon or Gulch
GabbsiWValley Range: s...c0.cseccce sacs <ccmdnes cueteeoteeceeeeeneeenee
Mayflower BallliMall 2.20024. ccscct spe secasschcecsenssesctecoenesse
IMUM Aces cot ceca sas ewese Socah os sveadwenens dead ceaeenowe aadeenceeaeceenecs

me SlATCKGOLPE ze casccssesecse cece sc Sawn ecee eee cises owone se so esietescateueese
MOLCHWESLEIN sas ee ae seseccces tenets canbe nce caceon ee ac conueee scetece cee decors
INBWO( GOS macteatnasuptoconcedacnoco CES RGECs ye aR SE nEE Ease hanna tncn aacaaaCee nas 70
WO NTONE CANYONS co gece cd oee eae os cae seesaw se ae oes ee ease cotesasae se 94
ah=Wte Range; Dun\ Glen Pass) .-.2.:....-:..:-.2s2--s<2sscsseesse: 100
IRE rSWING(COrs o-csgste ae: ccneee se ccscatescis sects sencsccessescsScseeatesse 70,71
ISAS THAN BO ree snc eee cea acne cucci ec dedsaecscuoses seecscuevecseeseeteee ses
Juniper Mine ...
OV CLOCK 2c ee nent cssessowees
Nevada Quicksilver Mine ... Js ae
NOSE! Creeks coc asseecw ase cece yo sees eeedsiaanosasc sansa cdsascacuvaaswesseses

WWESTEDMD S cois oac cane sees eceec Sens eee een Oa
New) Zealand <<... :csccces evevcscsshvcvesiecceectsccteuscnces one teeeenees
INE wtomii(ll9 87) aekccccecksstecne nes ccc ocousactosees nawecnsetehoncscesoesee sees
Newton [z7iprepillicecce.sceescesessseceseese=see ts
NewtoOn i Gait noite iecccccsccnsestos sence ec seeeeess oes
Nichols and Silberling (1977) .....................00+
NikolaitGreenstone) <...5cctsnececscecweseace seseeescncee cee eerste
Nizina’ Limestone Formation) <..c..ccs.0..2c-c0sseeceoneececeeters 73,88,93
Noli Bs ater ee ee a ek 93
INorthwAimeniGaus.sscces-t-seneressoceeeaesne nena 68-71,73,74,81,89,91,93
Notall re gionivs oo ioe. co.sccc st esceesnsasecsceaesaseectceetestsacecnsateootenere 68
“‘Nutzotin Mountains sequence” of Berg, Jones, and Richter (1972)
Jivaedesteay vacua ecteew resp ausbswrsssnace at eeergaaucered dadtweemecn tee seaacee ey 74
Ohios Woled ote. vases se asae sees scswencncseswawese ree neeeas eae esos 63
@ld ows OSA) sane csseswoeesveuetetaonomoosse se eee ee oe 69,70
Oldow, Bartel, and Gelber (1990) .................0..02205- 69-71,93,94
OTE GONE see eaee tease seneenoaee seotess oucedasswavencees 68,80,82,84,87-89,92
Crook Co
SUPICE area S sccc-oostaterec scenes aesos Be eetee eee terne sohete ee nae 72,94
AniBernard' Ranchvhouse) .crscsscssecsscccses-sec sesecessesesceees 94
Camp Creeker otek Seca a eae cdo ne ee coe eee eeecs 94
Paulina-Suplee moa). ceccs.cese-ceretedescseodeecscseetesea ceed ceece 94
-Paulinaitelephone ge-seeceaee se co reece cere ere eee reece eee 94
CASTER cco. ccndeacacwoeacde at were s casentunnceneucantearenaeeee tee eae eee 68
HelisiGany Ons nx. rcccseticcsevedene Saeecke eset cco ee qe
MONtHEASTEMM so. 2522201252 oss ov oodsn ee ccseew oe eee teste wae eeen tans Sena se ee 69
WoallowaiGo: <sicscsiciecea ieee ee ee ee eee 72
(Goon Greeks iicvccsswencss cess as cedoreses woke onc ause eee ee eae ane eae
Hurricane Greek ett; ete oo aero ee eee tach nese eee ee
Tm maha River secccccaccoccte cw sectie csc sencaeesacie cae seten sea stieestereeds
Little Granite Creek
McGraw Greek: wicccese-sceentceee ones
Snake River Canyon os
Wallowa: Mountains :::<.c...2.c2.n0n. csnsee cose acteotcsscscacuweences
@regon(State Wmiversity! ec <csseec-sccesccseocectascs cases sesscaesesereses
Orientospira Dagys, 1965 5
OSMANG! ZUSMAYEKEllAMee nw cth eos sosecedeee sone ie cteee coer seteT eee
Pacificiregion: eastern) <2... «...-sccsses-csesecencaserecenceeetee ser see cetse
Packard Washi Ronmationice-cc....c..csessconcaceecdscoserssecc stent see
Paleontological Research Institution, Ithaca, NY ......
Pamlico lithotectonic assemblage of Oldow (1984) ....
Papua NewiGuinea tac socsce ccccs cn cec scenes aes ce sus cavecusensesee eeeee
Paradisesterrane 2.22. -0crceceeee--ese-ce 67-69,70,71,81,82,84,91,92,94
parmata, Spondylospira ............... 10) 2282 66,68,79,83,84,100
panvan Vitimetulaes.cce-ccs Uno eeee 63,66,81,85,86,87,89,101
Pearson (UG Tine aoe camacceeeds decease seeesmmecenanne 87
‘pedicle tubules” of Yang and Xu (1966) 15
Pees, S. T., and Associates, Meadville, PA ................:.:00:000 63
IDEN DICXGPSCUAOSDONAVIOSPINGs aan cece ros ee ee eaten decieetre sasee eee tateee es ee
at ieee S ER Me Ne Nar ETN 10 ...... 63,66,68,69,79,80,84,89, 100
perplexa (aff.), Pseudospondylospira ..........6...4. LOM 2233 80,100
Petersburg A=3: 1:63,360 quadrangle: ...........5.<s.sssoesoeeeveveses- 92
Phenacozugmayerella, 1. gen. .........1.01e.c0eeseeeeeee 68,87,89,90,92
mimuncinata, N. Sp. ..........-+- M2 iecsoss 63,67,89,90,91,92,102
PiteShale Ae co. osc osecescsevcnne sh wens acecevaaes gat scassanevetonswseveseemeereemee ql
Port Alexander D-1 1:63,360 quadrangle ......................... 92,93
proto-Pacificiocean) DAaSINY-.cesc. sess ssccesvsceressorseceeseacsereeeet es 69,74
PseudolaballasMagys; NO Ancesescssssesesoceae ones necator eae ener 81,87
Pseudospondylospira, n. gen. ............6.5. 65,68,75,77,78,79,80,90
perpleXa;eSPs ...cescee~- 1h 63,66,68,69,79,80,84,89,100
alle PyperplexaNMuspy n..secsscssccoscetesess sete 10k 80,100
PStordeasHectoreal 857, 9 Mees ease saree eens ee rsa eee 86

ISPNS, [oaviceentots cowsnoeoewaterdeeaaoodencccnceusuece hewae cconseecaawonadeatees anes 64

108 BULLETIN 337

Pucara Group; Chambara Formation ...............-..--.------2+s 74,93
Py busiRormationsgresc0ss ccc cseecsc= oscsscescnncesenscoescnsae=cereseerren t= 73
Rail Cabin Argillite 72
Rangel Z. (1978) 81
REEGIS oise. cei tec feccorasccacieecocateesaseadeucenescaseeeek Uae cote aden eoueles 92
Reeside ental e(1197'8) oes ca socees coat cece Saree oe eee 2:
reesidei, Spondylospira ........ ... 81-84,86,99
1 tal (GIOLIS) \isssaesecdononasoccsase 66,69
RR AbdOCeraSISUCSSI:Z ONC esen- oa s see naecs honea secre ines teense eee 73
Raichardss Ro Woessteseerccstesctte ce <2: oeuseeec snsantere csececnunesteeam aces 92
Rachters(97.6) ies caceee cee ooo oe soe cns sa ce nee recone eS 73
Roberts Mountains Allochthon tes 71
Rocky#Mountainiiresiont ssc ..c-2-2c0c0ss2-02ecc2>ecsese-s-ecoeccsseesoscee= 71
RockysMountainviirenchits.ccc seeesa- noes eee seen ween ses heen = 93
Rosei@reekol'S quadrangle feec. -o. -- sec cans tore caanccece-c-aserevanee-ese 94
Seta rn errs re area eo oa eco oee costes noc seesssencaecens cacstcsctecsce-seees 94
Samisonrenala (U9 89) secacccerascoscaeoce vancetecence secs esos roe eee cee 69
San bormi(i9 GO) he sresrore nace coco cce coe ones Serer oa ee 71,83
Sanborns Acc b yo ccsssnceaasens canes onope Coenen soe oaeen dee rewe tacts anticsars 94
Sand Springs lithotectonic assemblage of Oldow (1984) ......... 70
Schathautli(8sil))c. ceae aden scossaceesocevees coarse veeee oeneeeoceeea 89,90,92
Schenk ESM cc. oic conc ase ones cies odoasacssateeessucteoceencessecedeoessweacsces 94
SchoonovermSequence ya. .isccaea-cancosnces tes cnes seco note oe sas aa seee econo 71
Seven Devils volcanics 94
Shimer/and'Shrocki(l944)i o-oo csecsmecenecs 64,81,86,99,100
Shoshone Meadows 15’ quadrangle .....................ss.ceeeeeeeeeee 94
Sierrav Neva Ga arGrncceecocesaseceacssacccsesscaedneasessdaceudiceeeeeesees
Silberlingi(US90) eo ses cssane cooneca=ceece- 5: sce ecna snot coe Ceon soe ee eee
Silberling and John (1989)
Silberlingverjaly (9 8i)lns.-cescceecaseacea-cseses=-eeses neste eee
Silberling and Roberts (1962)
Silberling and Tozer (1968) ............
Silberling and Wallace (1969)
SilberlingsJNG J), cc sccescasessenasecscetoeess 63,66,72,73,78,80,84,92-94
Sim MONS GAG ecw ecces cons cost evaene eaeeee oa eee enc ae te tee ene eee 94
Sirenites nanseni Zone [Zone LC3 of Tozer (1980)] ...... 66,82,93
Skwarko, Nicoll, and Campbell (1976) ...................ceeeseeeeeees 68
SmithrandFAeni(lS4)iiscscc. accewccccroncanncaseasessaancoeesase soseeeese 72
Snyder/and! Brueckner (983) <......2.c0s.c-cessersesesecuceatess eee -eaees 71
SOnOmasorOgenly, eee see eee es eee eee a ee Re 71
Sonoma Range lciquadrangles 220. 2s2e. 2.2 sseeee nce eceeeceoetg-ese-a <0 94
SOUtHVA Meri Can: recor ee reeecn: see ceeeee ce eeeececene ... 68,74,81,87
(GH iar rsre Panes Svcd ace eet oe cease cwaansus besneaescsesteneeoeese ee ntes 68
Perulyeseecdec sua sicewessascasset ie csacaessacseans 64,67,68,72,74,82,83,85
Atacocha District, Department of Pasco, Machican Mine area
Be Ri Fe Go ne ea aa gu a ace RUNGE TE RCP ATS COREE EES ROR 93
Carhuamayo 74
@erroydesPasco region y-cs-..c-scnccaccs-teawewncecceasettersesce os 70,74
Nimacaat ics: acciscesacseonscawcas soetedeudocnse tanec tecnesteeoeccesuoaes 74
Pasco 74
San Blas area 74
Yanamarca 74
dea: b.8. be ccc 64
fe 74,81,84,85
iyerella : 67,89,102
p. A; Zugmayerellat. ee 12-2 67,89,102

p. indet., Spiriferina(?)
peed (1977) <2 See heaves s- cones ooetoce eae eee eee
Speed, Elison, and Heck (1988)
SpinolepismatinaiDagys, 1974: x ..2 2. c..-<c.2- occas tee
Spirifera (Spiriferina) alia Hall and Whitfield, 1877
Spiriferina d’Orbigny, 1847

acrotamboensis Komen, 1937, -oesseseasceee enema nese 81,99
canavarica Tommasi var. robusta K6rner, 1937 ................ 81
koessenensis Zugmayer, 1882 81,87
Sp,ofmulmanny (917) Rees: -c-- ease 74,81,84,85
steinmanni Tilmann, 1917 .............. 74,81
suessi Winkler, W8590 5. 22.5. coc ecc. cc cnees oe sssnsoesssenence sone ee eneeeee
cf iS. suess? Winkler, 11859) .... ccc... <0 -ceed.ceses = oe
uncinata Schafhautl, 1851 .................-
Spiriferina(?) sp. indet. of K6rner, 1937
Spondylospira' Cooper; 1942) .........2..0<:-.-0+sso--0se-seaesseen eee eee
SER Oe SB Soke 63,64,66-68,70,73-75,78-80,8 1 ,82-87,89,93
acrotamboensis (Korner, 1937) <.:.<..<cs.-c-<:-<eeseaee eee 83,84
alia (Hall and Whitfield, 1877) .... 64,81,83,99,100
aff. S. alia Halland Whitfield! 2. ---- csc... --2s0--ceteseesceeeeeee 81
lewesensis (Lees; 1934) a.c.ccccecenssecesveecesccosos snot cee ter eee eee Ree eee
= 9,10 ...... 63,64,66,69,72,74,76,79,81,82-86,93,99, 100
cfiS: ewesensis\(ees,, U934)) «...ccccccccceeccsese- eee eee 81
parmata Hoover, 1983 ............. 10-45 66,68,79,83,84,100
reesidetiCOOpensy 1942 7 5.cc.conuacscassese Soe eeeeeeee eee 81-84,86,99
LFICOSEAN NSD ieeseeseeeo es Meee 63,66,69,74,81,83,84,85,101
Stanleyd(979)) re ee eee ee 64,68,72,86,90,92
StanleyG-. Di, Jie) 2osccsseccskesecesesaceacesoncce cos cteeere eee 63
Stanton), Tho We ,...cieoc0.6 scenes donccecseschecechecsectosteste tee ee 92
Star Peak Group, Cane Spring Formation ........................ 71,94
steinmannis Spiriferin@. ©. <2... <<c0.<4.csese0ccacnseceeese= ese eee 74,81
Stikineitermane occ soese tee caseceoeesceoe=ses =
Stone'and’ McWilliams) (1989)) <2ccccccccceceececcee- nese eee 74
StOnrS)Jy 5e.c0 050. ideedocenestanecnnsonedbohen neste aeeeun ee eee ee 92
Subcincularis; MONOUS: «<<. <.2os ce cn.ccscseaewscuensaseeta- eee eee 73,93
Subfamilies
Dagyspiriferinae, n-subfam:. :.......-.sce.cscncsesensosoece 63,77,79,80
Laballinae: Dagys; 1962. «......:...:-...s0s-sesq0dcaseaseeneereeeee 87
Spondylospirinae, n. subfam. ......................... 63,77,80,83,86
Thecocyrtellinae 75,77,80

subscripts [defined]
SHESSI, 0S DINIJENING 2. crscows se cenean-cceesonss

suessi (cf.), Spiriferina
Sun (LOB) ices. ccc aweecassewsncvanentnenssescacenccecee ate e tees
Sunrise Formation
Superfamily Spiriferinacea Davidson, 1884 ................. 67,75,81
Syracuse‘ University; Syracuse; NIY® <<2<.-..<c2cse-0cssuseccueeceeeeeace 63
Szekeleyand‘Grose'(!972)) <c..c.c.sss20ssee=snecsess snot sre eaee ee eee 74
Tantalus: Formation) 2.56.0... 5.5 oe eesceck cscs ceeset eee 73
Taylor:et al. (U983) cic de ct hack deccdeas savave soso eee 70
Paylor D. |G: .20cc02cideec bev ccvews tee a es ee 72
“Methyan*;'Species) .2.c.<c-<-sescsevesnerees .... 63,68,69
Meth ys BPs ee eee 66,68
CASEIN 5 505 cccsiveececcesedes desea ce oleePave aces tao ene 68
WESIEED . 55 Ss ccs deesvectcecua es cctiva Sac cdeet pee eer 68
Thecocyriella Bittner; 1892-28. e. se. 67,75
Thecocyrtelloidea Yang and Xu, 1966 67,75
Thorman etal: (W990) ect e ete ene n ee 71
Mibet ences sscce ose saccsscoe von ee aero 68,87
Tilmann (1917) x 64,74,81,84,85
Tonopah: 1°‘quadrangle \.. 30. :<.2-cccsnen acc soctvaston sooseesseeeeee eae 94
Dozer(UGS8)) .sscscccsaceiovenccscdeccocs vaccnns vane sWomees <SaRe eee eOe 73,93
Tozer (1962) sess cczaccci sees eeseds 2 ocdesnsks oocuse swans eed ee Ee 81
Mozer (980) sereacceecess-s eee 66-68,78,79-82,84-87,89,91,92-94
LOzZEr' (982) coccacasecccewevhsccscaueaevecesasessseeeeec aces ecces eee 71
Ozer (UGB) csc cove cecsocncensewsdovcseoss oesetaewcsen ease 70
TOzZernE: TDi Sasciccsec bcs ccconacsenc scorers ee 63,93

Trachyceras aonoides Zone [included in Zone LC1 of Tozer (1980)]
sabes si eecheeedeses teat ee, Slee ee 66,82,93

LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 109

Trachyceras desatoyense Zone [included in Zone LC1 of Tozer (1980)]

Pa Pn Me Sock oe ale sot nses og cahwics podeenayaceutevece arts seeeess or ea 66,82,93
tricosta, Spondylospira .... V1 ...... 63,66,69,74,81,83,84,85,101
Tiropiceltitesich Ti: COMMDIANUS) 2. .c..c00ccescc0seenseasesesscecscesseos- 93
Tropites dilleri Zone [Zone UC1 of Tozer (1980)] ..................22.5

Me eet a ne sta sea ccresheeceeesstass 66-68 ,73,79-82,89,9 1-94
Tropites subbullatus Zone [included in Zone UC2 of Tozer (1980)]

PR eh Orns: cols nce Na csgeeeets 66,67,72,73,80—-82,88,89,91-94
Tropites welleri Zone [included in Zone UC2 of Tozer (1980)] ......

Pees era ncyNecccas bec ssedeas suka 66,67,72,73,80-82,88,89,9 1-94
WiUITKeEVAMONtDWeStEM!, +.05.52-csc0c ce vceecace ster anestercceicdecens<naecnenese 68
WU GHC SMID seers = Sciocu shan secss sp eetusssewetebeeredeued oe 63
MhyaughtOnl Group ce. s-c--0-eeee oe ssactee es sess vc vocnareceeos ea aceaee seen ees 93
uncinata,

SS DUME) CMI itrowr ccc nc cose tee oie. seen ons eeetae eee ae ccc swe een ee aucseets eee 90
EUS IVILYV EVOL Oe settasnasinc reg cows sareceetce ten sccesenenewesse 64,89,90,92
University of Montana, Missoula, MT .......... Jeeeasaseescswaewies 63
“Upper Triassic limestone” of Richter (1976) 73,74

USGS [United States Geological Survey, Paleontology and Stratig-

raphy Branch, Washington, DC] ...................02000+ 63,65,74
USGS localities
DS 09K(Mocalityill)| sces.cccsscecceesssececeeeesecsesesaceeeses: 71,89,92,102
DSA (Wocality 2)! sicw.tercescee save sscseresactiascesveseseesee esters 71,84,92
DAA Gi (ocality 3)! cec-ssscscsare-cnessccsnecesesse- 71,88,89,92,101,102
DAGON Meacality co) ives ssc .ce sec netes sce se ssesescecstesees oveeeeeess 71,81,92
NOS48i(eocality6))2s/-ce-.cneeezceccrnec.sccteseenere-cees 67,73,81,91,92
NA O GN(Wocality 97) \-sc-ce-222-ecseree ste ssceesenesaauecesecastees oes 73,79,92
IUISHAN (eocality28) a-scsene0< ese os occ cee ace sccetcceeee ec ee 70,84,85,92
NB3S54i(Hocality lO) oeescccc.csce- cee sescteoccosusnceosesccees sews 70,81,92
SSS Ol(ocality 2) ec. 2tec.-coseteceescceseesencersons 70,72,89,92,102
N62 Gi (Wocality 313) sic. <ccc sees see eee sw ssee=escoseresesees 73,81,88,92
17424 (Locality 15) = ‘tthe Lewiston locality” ......................4.
eae ee onto Patan sites beet oe ee 72,81-83,86,87,92,93,99,101
USGS M [United States Geological Survey, Paleontology and Stra-
tigraphy, Branch; Menlo Park,’GA]| .............:----s-------- 65,74
USGS M localities
MO Gi (locality ll8) ieee eve acces cacccc.cdenscsetesccns coseses 70,79,93,94
M1708 (Locality 20) .... 73,88,93,101,102
IML 4 Oi eocality: 2) e.cc ice essa se sees seseoncoves-sceneses 72,82,88,93
INAS O Gi (ocalitys2.2)i 2) ssccescst cove ce ssedeces este aeceseore 73,80,93,100
M1911 (Locality 23) .... 73,77,78,80,93,99
MINGUS (ocality224)i 2s. cess cesdcecss steve sy ¢scenccs ae-eseete 73,80,93
M1919 (Locality 25) .... Paatos ts shochu eeancn tee secenes 73,80,93
NED 135i (ocality,.26)) tices csnccescesseenetescctsseseeseseeee- 66,73,82,93
NZS 6i(ocality:2/7)) -ss.c.-.:.s+-csnsc-tceveennece sess: 73,80,82,93,100
M2672) (ocality 28)) (ci s.c-+.cx.c-2-0ese0=s 72,80,84,88,93, 100-102
USGS [United State Geological Survey, Denver, CO] ........... 63
USNM [United States National Museum of Natural History, Smith-
sonian Institution, Washington, DC] .....................2:..:0ee0eee
De cedece seduce dies siecaee deseseees 63,64,74,77,78,80-83,85-89,99-102
WESaNationallResearch:: Council! e2.. 22-2---ces-e5-ceeceeceneeceee ree sees 63
W.ES:S.R..,
CaTicasusiMOountalnspeeeceretseseesees ere sees ce ee ree ne 68,87
MOTtHEASTETD ureter aces a reses teens ees arte re ner ner one oraear Meeeeee 68,87
MOLEMEASTETMIM OSU ore set a seceeer tects e tee terea ter ese nee ta renrace ae 68,70
IKoryak Mountain Range: <2. .02c.s-.-s:scseccesscesterese«ceesess 68,81
ORM OTS errs seers sete eres eo evateteunecuaenbuns evivinctacede eevee 90
SIDED A aor s oon Fac sean les oie fee vecus sesecsssevecsscenssse sccetesselecsen sts 68
Paya Celine pares mre creas cenc tance. eee canteen ene a aoe 74
Waldes,. J. ..:.-::.: aes 63
Walliers(Q19 6:7) ee ssa acsccce soe sene nee fe aeene enc ena toopen sere ane case sseene eee 93

Walls eresete ple, ercecs soncske sete nes se teseoreresatenswnanctessestmascscWareseseue ws 93

Wanider Vioovel ai N(19 80) se rnacassrencasereretscessstaacaeceronanencecees 73
Wigrass, Ib. Wis ccsstsicesecnscseses 94
Vitimetulaim S6Ds <c2a.-2s.¢2se2eenenarese= 68,75,80,81,83,85,86,87,90

PArVas Ms SP ies socsseeeeeerereee 1 eerie 63,66,81,85,86,87,89,101
Vokes:and: Haas! (1944). 2.5 och ccccetecces oes. scowscseeeesssceses 64,74,81
Waageni(l'883S)):ceiccac aoc custece seeecusececacse vacceasscesserste ssesdenteans 75
Wales Swansea s.2-c e045 coe coos ener ne ae eee en ae Sac nen ted seen eeeee 63
Walker Lake terrane of Silberling et a/. (1987) ...............0..... 70
WaallacéuR® Es sA.sctis -<-css dss wees oa tam ace ras evoees aneataen tees ne tnen ats 93
WallowarBatholithy fac seces cocese ses nsacee soseedes econmenevs sacnaseseunseee 72
Wallowa'terrane ....-...-.--...--- 68,69,72,80-82,84,86,88,89,92-94
Wangi(i955)) chsteccces scteact ceecses sec noe ses osu aes sanconenectcctetbecsees 87
Weashburne3G: ssc. ceccsecbes ccssadeaeeeessentusoraenovasseiaresaacvestsecsa: 92
Western¢Pacific:sregion® 2.2.2.-c-c-2e0-.52-cssee ooo: cannes eee eeceseeases 68
Wheeler HE s 5 cv. ccd scaic seuss sacsuscossssz se ncdsbac tess veasasssaesususss 94
Williams: JaS° =.-:--:- ee ae ae See Peron
Wankler.((US59) i.e sie. nse o.saccorsicacswaseesedonescetOusceard goceusscocees 83
Wrangelliaitemmane=---..c..-c-coecseeceseee- oe 68,69,73,74,81,88,92,93
Yangrand Xiri(l9 66) cose. ccaneencessccessceceeesssselevenscdsuctescaaans 67,75
VANOSDIFAMD ABYSS WOT = sosccsenesecee-caceecedeeeorasceacontesecest tte 89,90
Yole:and'Irvingi(1980)! set ere sec qoanesocetoneacerteceeecteeesacresect 74
yuellangpingica, Zugmayerellacce..-sessccenes-seceecersen-seeseenes cate 89
Zlambach: bed Si ccc. 22 .cceas sass owaae sc aoacdesscaseccenceesoeskaceneoeesanaees 93

Zones of Tozer (1980)
LC1 [Trachyceras aonoides Zone; Trachyceras desatoyense Zone]

Ae seesk See beers Senseee Sees Sma aBECT ooC ane OC ROSALES 66,82,93
LC2 [Austrotrachyceras austriacum Zone] ..............--- 66,82,93
G3 )\[Sirenitesinanseni-Zone]\..<.---s22-2-eeteseceens eens ees 66,82,93

LNol [Guembelites jandianus Zone; Mojsisovicsites kerri Zone]
66,67,78-82,84,88,89,9 1-94
LNo2 [Malayites dawsoni Zone, Malayites paulckei Zone] ........
eee OE oe 66,67,80-82,84,85,89,9 1-94
LNo3 [Juvavites magnus Zone] .... 66,67,73,80-82,84,85,9 1-94
MNol [Cyrtopleurites bicrenatus Zone, Drepanites rutherfordi Zone]

BRAS RE eee ere Si a eee EEE 66,80-82,84,9 1-94
MNo2 [Himavatites columbianus Zone] ...............2..1.00-00-00++0++

Be ease eee race cere st arere see ceee 66,67,80-82,84,9 1-94
UC1 [Tropites dilleri Zone] ............ 66-68,73,79-82,89,9 1-94
UC2 [Tropites subbullatus Zone; Tropites welleri Zone] ............
Cee ae ES COR Re So cena -eeracceLn co 66,67,72,73,80-82,88,89,9 1-94
WG3[Klamathitesimacrolobatus'Zone] |c.-:-.2-ceecereceeceneseaseoset eee
PR Rrra saree More REE A Sr eeaa te ce ee 66,67,80-82,88,89,9 1-94
UNol [Gnomohalorites cordilleranus Zone] .................06..000004-
SAR MTS ORNS SaaS cea feed a Ona eee ah ae 66,68,80-8 2,84 ,86,92-94
UNo2 [Cochloceras amoenum Zone] ..... 66,81,82,84,86,92-94
UNo3 [Choristoceras crickmayi Zone] .............00.:00c000ceeeeeeeeeees
Sears eR SEGA SEAT nO EAD BORER PEATE 66,68,81,82,84,86,92-94
Aupmayenn(ss2)i sec ses sees ose teceecse seer Mean eee 81,87,89
Zugmayerella Dagys, 1963 ......... 63,64,66-68,70,81,85,87,88,89
americana, N. sp. .........-- sh ES ee 67,86,87,88,89,101,102
CureaMagyss V9GS: -cscsecscccds -seanscesuesceceee ceca vases ee swec eae seeseees 89
REMIDYrAMIAG SUDA OSI esccse sete nas esate see eeee neta cere oe
inaequiplicata Dagys, 1965 89
WPICINQLO Ne. 5 Foo sates ssinds vedic sus vesdauin oo ceaieeeaeatees iceaeeees nace Al)
Koessenensis\(ZUgMayers USS2)io.cscs.nececerseccecs coceseesscecceseees 89
OSMana (Bittner 11902) Mreseccscssesceceesecectresesec tar cnoaeeeeceteee 89
SD eat aca wenn een eae aaa Soa nse Seen 1 eee 67,89,102
uncinata (Schafhautl, 1851) ..................... 64,89,90,92
yueliangpingica Ching and Feng, 1977 .......................00006- 89
ZUSMAVEFElIATISP Agree eneenereetn ee eee 1 Pere 67,89,102

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